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Full text of "Mammals of the Soviet Union"

Mammals 
of the 



oviet 




.иМЕП 

Fan la 



VG. Heptoer 



N. E Naumov 



r"^ 



■L-lr-?^^ 



:•■'■' . o.^ 



The present book, of the series Mam- 
mals of the Soviet Union, Sirenia and 
Carnivora, is the fourth volume to be 
pubHshed in EngHsh. Previously Volume 
I — Artiodactyla and Perissodactyla, 
Volume II, part 2 — Carnivora (Hyenas 
and Cats) and Volume II, part 3 — 
Pinnipeds and Toothed Whales were 
published in 1988, 1992, and 1996 
respectively. Sea Cows, Wolves and 
Bears constituted the second in the 
series to appear. 

The original Russian book is so 
large (982 pp.) that its publication in the 
English translation format would have 
resulted in a book of about 1400 pages, 
which was considered too unwieldy. 
Therefore, it was decided to publish 
Volume 11, Part 1 — Sirenia and Car- 
nivora, in two roughly equal -parts. The 
first (Part la) consists of the Steller's sea 
cow, the now-extinct marine mammal 
once found only around the Commander 
Islands in the western Bering Sea, and 
the Russian species of the wolf and bear 
families. The second half (Part lb) will 
comprise the various members of the 
weasel family, plus the introduced 
American raccoon. As indicated in the 
foreword to the English edition of 
Volume I, the senior author and editor 
of the series. Dr. Vladimir Georgievich 
Heptner, died in 1975, and did not 
complete the projected series. However, 
work continues, and Russian volumes on 
baleen whales, lagomoфhs and dipodid 
rodents are now published. 

A final point: since the appearance 
of Volume II, part 2, the Soviet Union 
has disappeared. However, the series was 
written in the context of the former 
"union of republics," and it would be 
confusing to re-edit the text to conform 
to the current political reality. 



Mammals of the Soviet Union 

Volume II 
Part la 



MAMMALS OF THE SOVIET UNION 

In Three Volumes 



Edited by 
V.G. Heptner and N.P. Naumov 



Vysshaya Shkola Publishers 
Moscow, 1967 



Mammals of the 
Soviet Union 

Volume II, Part la 

SIRENIA AND CARNIVORA 
(Sea Cows; Wolves and Bears) 



V.G. Heptner, N.P. Naumov, P.B. Yurgenson 
A.A. Sludskii, A.F. Chirkova and A.G. Bannikov 



Illustrators 
A.N. Komarov and N.N. Kondakov 



Scientific Editor 
Robert S. Hoffmann 




Smithsonian Insitution Libraries 

and 

The National Science Foundation 

Washington, D.C. 

1998 



Mlekopitayushchie Sovetskogo Soyuza 

In Three Volumes 

V.G. Heptner and N.P. Naumov, editors 

Vysshaya Shkola Publishers 
Moscow, 1967 

Translators: Bolos Abdul Malek Botros, Hamed Tantawi, Hosni Ibrahim 

Youssef, Ali Abdul Moneim Moussa 
General Editor: Nawal Mehallawi, Al-Ahram Center for Scientific 

Translations, Cairo, Egypt 

© 1998 Amerind Publishing Co. Pvt. Ltd., New Delhi 

Library of Congress Cataloging-in-Publication Data 

(Revised for volume 2, pt. la) 

Geptner, V.G. (Vladimir Georgievich), 1901-1975. 
Mammals of the Soviet Union. 

Translation of: Mlekopitaiuschie Sovetskogo Soiuza. 

Bibliography: v. 2, pt. la, p. 

Vol. 2, pt. lb has index. 

Supt. of Docs, no.: Si 1.2: Ar 7/2 

Contents: v. 1. Artiodactyla and Perissodactyla — 
V.2. — pt. la. Sirenia and Carnivora (Sea Cows ; Wolves and Bears) 
v. 2. — pt. 2. Carnivora (Hyaenas and cats). 

1. Mammals — Soviet Union. I. Nasimovich, A. A. 
II. Bannikov, Andrei Grigor'evich. III. Hoffmann, Robert S. 
IV. Title. 
QL728.S65G4713 1988 599.0947 85-600144 

Published for the Smithsonian Institution Libraries 

by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110 001 



Printed in India at Baba Barkha Nath Printers, New Delhi 



Foreword to the English Edition 



The Smithsonian Institution Libraries, in cooperation with the 
National Science Foundation, has sponsored the translation into 
English of this and hundreds of other scientific and scholarly studies 
since I960. The program, funded with Special Foreign Currency under 
the provisions of Public Law 480, represents an investment in the 
dissemination of knowledge to which the Smithsonian Institution is 
dedicated. 

The present book of the series Mammals of the Soviet Union, Sirenia 
and Carnivora, is the fourth volume to be published in English. The 
project was initiated in 1975 as my contribution to joint U.S. — U.S.S.R. 
studies on Holarctic mammals, sponsored by the Environmental Agree- 
ment between the two countries. Previously, Volume I, Artiodactyla 
and Perissodactyla, Volume II, part 2, Carnivora {Hyenas and Cats) 
and Volume II, part 3, Pinnipeds and Toothed Whales were published 
in 1988, 1992, and 1996 respectively. 

Sea Cows, Wolves and Bears constituted the second in the series 
to appear. The fact that this volume was not published in English 
translation in the proper sequence is due to the fact that the translation 
was performed under a different contract from that supervised by 
Amerind Publishing Company, New Delhi, India. Moreover, the origi- 
nal Russian book is so large (982 pp.) that its publication in the 
English translation format would have resulted in a book of about 
1400 pages, which we considered too unwieldy. Therefore, it was 
decided to publish Volume II, Part 1, Sirenia and Carnivora, in two 
roughly equal parts. The first (Part la) consists of the Steller's sea 
cow, the now-extinct marine mammal once found only around the 
Commander Islands in the western Bering Sea, and the Russian spe- 
cies of the wolf and bear families. The second half (Part lb) will 
comprise the various members of the weasel family, plus the intro- 
duced American raccoon. 

As indicated in the foreword to the English edition of Volume I, 
the senior author and editor of the series. Dr. Vladimir Georgievich 



VI 

Heptner, died in 1975, and did not complete the projected series. 
However, work continues, and Russian volumes on baleen whales, 
lagomoфhs and dipodid rodents are now published. It is hoped that 
English translations of these later volumes will follow more promptly 
than has been true of the first volumes of the series. 

English readers interested in Dr. Heptner' s contributions as a mam- 
malogist should refer to the foreword to the English edition of Volume 
I. Conventions used in rendering geographic names, first stated there, 
are reprinted here for the convenience of the reader. Geographic names 
are generally transliterated directly, but a few exceptions were permit- 
ted (e.g. Moscow instead of Moskva, translation rather than 
transliteration of certain modifiers of place names, such as Northern, 
rather than Sevemaya, Dvina). Soviet administrative units are numer- 
ous, and the following equivalents were employed in translation: krai, 
territory; oblasf, district; raion, region; guberniya ( archaic), gover- 
nance, uyezd, county. Also, in the original Russian text, rivers, mountain 
ranges, and cities are often not explicitly identified, the Soviet reader 
being presumed sufficiently familiar with the geography of the coun- 
try to be able to understand from the context of the sentence what sort 
of place is referred to. Complicating the matter is the lack of articles 
as parts of speech in Russian. To assist the English reader, I have 
adopted the following conventions: if a river is referred to, an article 
precedes it; if a mountain range is referred to, it is translated as a 
plural; if a city is referred to, it is singular, and lacks the article. 
Examples are: the Ural (river); the Urals (mountains); Ural'sk (city). 
Geographic place names are also inflected in Russian, and these have 
been simplified by omitting transliteration of the inflected ending. For 
example, the Russian phrase v Yaroslavskoi i Kostromskoi oblastyakh 
is translated "in the Yaroslavl and Kostroma districts." In cases where 
the nominative form of the place name has an -sk ending this is, 
however, transliterated (e.g., Omsk); when a Russian "soft sign" is 
employed in a place name, this is transliterated as an apostrophe (e.g. 
Khar'kov); the "hard sign" is rendered as a double apostrophe. Be- 
cause of the large number of place names in this volume, it was not 
possible to verify all of them, and some inconsistencies are likely to 
occur. I would appreciate it if readers would bring any errors they 
may notice to my attention. 

In Volume II, part 1, I have also attempted to improve certain 
aspects of the translation in order to reflect more faithfully the original 
Russian text. For example, what was translated as "taxonomy" in 



vu 



volume I is now in some places rendered as "systematic position;" 
what was "economic importance" is now "practical significance," etc. 

One further point of confusion not apparent to me when Volume 
I was translated also required clarification; that is the English trans- 
hteration of the senior author's surname. This begins with the fourth 
letter of the Cyrillic alphabet, which usually has a "G" sound in Rus- 
sian. However, the surname was originally German (as indeed was 
mine) and in the original German began with the letter "H" of the 
Latin alphabet. Since Cyrillic has no equivalent of "H" this is usually 
transliterated into "G" in Russian. However, I know from conversa- 
tions with him that Dr. Heptner preferred to use the original Germanic 
form of his surname rather than the transliterated version, which is 
rendered as Geptner. The rules of transliteration employed by the 
Library of Congress do not permit such flexibility, and the attentive 
reader may notice that Library of Congress cataloging employs the 
latter. 

A final point: since the appearance of Volume II, part 2, the 
Soviet Union has disappeared. However, the series was written in the 
context of that former "union of republics," and it would be confusing 
to re-edit the text to conform to the current political reality. 



Robert S. Hoffmann 
National Museum of Natural History 
Smithsonian Institution 
Washington, D.C. 



Foreword 



The second volume of the "Mammals of the Soviet Union" is devoted 
to descriptions of the terrestrial Carnivores (Order Camivora), and the 
sea cows (Order Sirenia), consisting in our fauna of only one recently 
extinct species. There is a great deal of information in our country 
about carnivorous mammals, which include the most important fiir 
animals. In the last 10 years they [furbearers] have been studied in 
especially great detail. Various unpublished data of importance avail- 
able to the authors were included in the descriptions. Therefore, the 
size of this book is considerable. 

All species in the indigenous families of the Order Camivora are 
described with the exception of the cats (Felidae) and hyaenas 
(Hyaenidae), i.e. the suborder Feloidae. Descriptions of these excluded 
families will follow in the next volume. In addition, the book includes 
two acclimatized American species of carnivores — raccoon and mink. 
-The striped skunk, as is known, failed to be acclimatized in our 
country and is therefore not included. 

The classification of this order follows Simpson (1945). Simpson's 
orders themselves are, as is known, conventional. Only one exception 
is admitted: pinnipeds, contrary to the opinion of Simpson and some 
others, are considered as an independent order. The arrangement of 
orders in the book follows the sequence from most specialized to the 
most primitive, that is to say, from up to down. Within the orders in 
this volume — as in volume I — the taxa and species are arranged, 
according to their specialization. In the sequence of orders described, 
however, one other deviation was made: carnivores (Camivora), and 
not pinnipeds (Pinnipedia) were placed after the sea cows, in order 
that pinnipeds and whales could be included in one volume. For ori- 
entation to the actual relationships of orders, a macrosystemic scheme 
of the class is given [pg. xix]. 

All the characteristics of a group are presented in a standard for- 
mat and deviation was permitted in only a few special cases. Group 
characteristics, given in an extremely condensed form, are applicable 



not only to the species of our fauna but to the entire group. Principal 
attention is paid to the distribution and systematic composition of 
groups, and the phylogenetic relationships of our forms within the 
group. Paleontological information is very condensed, mainly accord- 
ing to Simpson (1945), but partly from Thenius and Hofer (1960), 
"Fundamentals of Paleontology" edited by Yu.A. Orlov (1962), and 
various other sources. 

In the species descriptions, attention was mainly concentrated on 
geographic distribution and biology. Ranges are broadly presented, 
based on available data, and only limits are described. Habitats within 
these ranges are not mentioned, and only the most important marginal 
localities are given. Historical changes in geographic ranges are traced, 
and as much as possible the description is based on the "reconstructed" 
range of the species, in depth for centuries of "historical" time. This 
is, therefore, a suitable opportunity to establish a more or less actual 
geographic area. A knowledge of this is obligatory for solving theo- 
retical questions of zoogeography and for practical work with the 
fauna'. 

Ranges beyond the boundaries of our country — ^including those 
"reconstructed" — ^follow numerous works, but chiefly for critical veri- 
fication, information provided by Seton-Thompson (1909—1910), Miller 
(1912), Sowerby (1923), Anthony (1928), Aharoni (1930), 
Nezabitowski (1934), Shortridge (1934), Phillips (1935), Heim-de- 
Balsac (1936), Allen (1938-1940, 1939, 1942), Pocock (1939, 1941), 
Cabrera and Yepes (1940), Hamilton (1943), Нафег (1945), Simpson 
(1945), Anderson (1946), Carter, Hill and Tate (1946), Prater (1947), 
Tate (1947), Troughton (1948), Enar (1949)*, Calinescu (1951), Rob- 
erts (1951), Ellerman and Morrison-Scott (1951), Morrison- Scott and 
Hayman (1953), Bannikov (1954), Laurie and Hill (1954), Miller and 
Kellog (1955), Siivonen (1956), Haltenorth and Trense (1956) Cabrera 
(1958, 1960), Van den Brink (1958), Frechkop (1958), Markov (1^59), 
Misorine (1959), Hall and Kelson (1959), Hatt (1959), Dulic and Tortric 
(1960), Burt and Grossenheider (1962), Burton (1962), Cockrum (1962) 
and many others. Individual references to the above-mentioned au- 
thors were not given in the text descriptions of ranges external to the 
boundaries of the USSR. North American ranges were based in nearly 

' More detailed information about reconstructed geographic ranges is available in 
the work of V.G. Heptner. Dynamics of geographic ranges of some ungulates and 
human factors. Sb. "Geographical Questions", 1959. [Dinamika areala nekotorykh 
kopytnykh i antropicheskii factor. Sb. "Voprosy Geographii," 1959.] 

* Not in Lit. Cit.— Sci. Ed. 



XI 

all cases on excellent maps in Hall and Kelson (1959), whose opin- 
ions differed completely, in some cases, from the description of others. 
All maps of geographic ranges are original, prepared by V.G. Heptner. 

Unfortunately, we have not succeeded in achieving uniformity 
among different authors in describing the biology of species. How- 
ever, the general format is identical to that encountered in the first 
volume; only the description of the extinct species— the sea cow — 
differs. Concerning the latter, instead of paraphrasing the data from 
published literature, a special translation was made of the original 
work of Steller — the only naturalist to have observed this animal and 
studied its anatomy. Until now, i.e. for more than 200 years, Steller' s 
text has not appeared in Russian. S. Krasheninnikov (1786), who had 
Steller' s manuscripts, published a translation of only certain parts of 
this work. Owing to the language in which the work of Steller was 
published (Latin and old German of the early 18th century), this 
material was not accessible to most zoologists, as a resuh of which 
some disagreements arose in our day. Knowledge and iconography of 
the species was poor.* 

The moфhological characteristics of the species have been pre- 
sented in completely identical, though short, form as far as sufficiently 
detailed descriptions of the series of taxa and forms. Geographic varia- 
tion of species within the USSR has, with few exceptions, been revised 
or critically reevaluated. Variation in those parts of geographic ranges 
lying beyound the boundaries of the USSR follow the most recent 
available data, but were not critically reviewed; the described forms 
are listed and general notes provided. 

All synonymys are re-examined and given in an extremely limited 
scope according to the following principles: only species synonyms 
are included. Generic synonyms are mentioned only if the type of the 
genus is a species represented in our fauna, or if the generic reference 
has significance in that it sometimes has been applied (or previously 
applied), or may be applied to our species. All references are included 
in species synonyms in the following cases: 1) if the name was ap- , 
plied to a specimen of a species in our territories; 2) if the name was 
applied to a specimen of a species taken elsewhere, but is significant 
in that it is applied or has been applied but is now used for a Soviet 
form; 3) if the name was given to a specimen from another territory 
and not applied to our form, but may have significance for some 

* Steller's text is now known to be inaccurate in certain respects — Sci. Ed. 



Xll 



Soviet forms; 4) the names applied to type species are always pre- 
sented regardless of their type locality within or outside of the USSR. 
Actual names in the species synonymies are not alloted, but are placed 
in the accounts of subspecies inhabiting our country, and synonymies 
are also given in relation to the particular subspecies. Synonymies of 
subspecies distributed outside the USSR are not given. Synonymies of 
categories above the genus are not mentioned, nor are references to 
the descriptions and names of these higher orders which, if not stated 
otherwise, are given by Simpson (1945). 

Names applied to domestic forms, following the rationale and 
usage of the first volume, are not used for wild species; rather the 
name assigned the wild taxon is employed for the species as a whole. 

The total number of mammalian species approximates 3500, and 
the number of species of our fauna is about 300 (Heptner, 1956). 

Each author's contribution in the compilation of this volume was 
as follows: V.G. Heptner wrote all the general characters of the major 
taxa (order, family, genus), the entire sections concerning the sea cow 
(including translation of Steller's text) and the Afghan fox, and all 
tabular material. He worked out the synonymy of all species and also 
wrote the sections dealing with the "Description", "Systematic Posi- 
tion", "Geographic Distribution", and "Geographic Variation" for all 
species described in this volume, and also prepared all maps of geo- 
graphic range presented in the volume. P.R. Yurgenson wrote the 
section on "Biology" and "Practical Significance" for the three spe- 
cies of bears, all species of the mustelid family, and the American 
raccoon. A. A. Sludskii wrote the same sections concerning the corsac 
fox, jackal and red wolf [dhole] whereas the section on "Biology" and 
"Practical Significance" for red fox and arctic fox were contributed by 
A.F. Chirkova; N.P. Naumov wrote the same sections for the gray 
wolf, and A.G. Bannikov, for the Ussuri raccoon (raccoon dog). 
Authorship of the separate parts is indicated by bracketed initials at 
the end of the corresponding sections. Integration of the [above] sys- 
tem, species arrangement, their limits, number and limits of subspecies 
as well as all the synonymies are the responsibility of V.G. Heptner. 

The literature up to the period 1961-1962 was more or less com- 
pletely employed. Some works which appeared after that period were 
not referred to, especially the information provided by I.M. Gromov, 
A.A. Gureev, G.A. Novikov, I.I. Sokolov, P.P. Strelkov, and K.K. 
Chapskii ("Mammal fauna of USSR" 1963). We have succeeded in 
only individual cases in making use of data from papers published 



XIU 



after this period. Literature references are usually given parentheti- 
cally by surname and year, initials being given only when more than 
one author has the same surname. References to author's unpublished 
data, personal communications, or letters are given without reference 
to year, but the author's initials are given to distinguish them from 
published citations. 

The authors have, of course, included their unpublished personal 
observations in this work, and this is sometimes referred to as noted 
above. In addition much unpublished material of different types was 
kindly supplied to the authors by G.F. Bromlei (Vladivostok), N.K. 
Vereshchagin (Leningrad), N.N. Vorontsov (Moscow), S.V. Kirikov 
(Moscow), V.V. Kozlov (Krasnoyarsk), A.P. Komeev (Kiev), V.P. 
Kostin (Tashkent), B.A. Kuznetsov (Moscow), N.I. Larina (Saratov) 
Kh. I. Ling (Tartu), R.N. Meklenburtsev (Tashkent), L.G. Morozova 
- Turova (Moscow), V.N. Nadeev (Irkutsk), T.A. Pavlenko (Tashkent), 
V.Ya. Parovshchikov (deceased) (Arkhangel'sk), V.N. Pavlinin 
(Sverdlovsk) V.A. Popov (Kazan), S.K. Priklonskii (Oka preserve), 
R. Rausch (USA), С Repenning (USA), N.N. Rukovskii (Moscow), 
N.V. Rakov (Saransk), V.E. Sokolov (Moscow), V.N. Skalon (Alma- 
Ata), S.U. Stroganov (deceased) (Novosibirsk), N.D. Sysoev 
(Vladimir), K.A. Tatarinov (L'vov), V.P. Teplov (deceased) (Oka 
preserve), V.V. Timofeev (Irkutsk), S.M. Uspenskii (Moscow), A.V. 
Fedoseev (Bryansk), K.K. Chapskii (Leningrad), V.D. Shamykin (Mos- 
cow), E.I. Shereshevskii (Moscow), E.I. Shcherbina (Ashkhabad), as 
well as additional authors especially mentioned in the text. 

Useful information concerning the distribution of the different 
species was also kindly supplied by workers of all the district hunting 
inspection centers. These were Starodubchenko (Kiev), Lebedev 
(Volgograd), Mikhailov (Saratov), Zakharov (Murmansk), Gusev 
(Rostov-on-Don), Khmelevskii (Orenburg), Sukhikh (Belgorod), Ivanov 
(Yakutsk), Kuz'min and Fofanov (Novosibirsk), Bel'skii (Vladivostok), 
Anashkin (Ulan-Ude), Samsonov (Blagoveshchensk), and Sysoev 
(Khabarovsk). 

Figures for this volume were assembled by V.G. Heptner. Animal 
illustrations were prepared by our famous "animalists", A.N. Komarov 
and N.N. Kondakov. Line drawing of all of the animal species discribed 
as well as five of the color plates, were made by A.N. Komarov. 
Three plates — ^weasel, red fox and sea cow were done by N.N. 
Kondakov. The painting of the Steller's cow is considered a trial 
reconstruction of the species — a collaboration between N.N. Kondakov 
and V.G. Heptner. The drawing of tracks were, with few exceptions. 



XIV 



taken from Prof. A. N. Formozov's books based on his original sketches 
from nature. 

The skull drawings were done by V.N. Lyakhov based on mate- 
rials in the Zoological Institute of the Academy of Science, under the 
supervision of Prof. G.A. Novikov, and were previously published in 
his book (1956). The original drawings are in the possession of the 
Zoological Institute of the Academy of Science; thanks to the courtesy 
of Prof. O.A. Skarlato and Prof. A. A. Strelkov, the Institute has granted 
permission for their use in this book. 

Photographs and other illustrations related to the biological sec- 
tions of the volume were contributed by various persons, but were 
mainly taken from the archive of Prof. V.G. Heptner (especially those 
of mustelid and bear families). A significant portion was provided by 
A.A. Sludskii (Alma-Ata), Yu.K. Gorelov (Badkhyz preserve), S.V. 
Marakov (Kirov) and A.G. Pankrat'ev ("Kedrovaya pad" preserve). 
Those supplying other photographs are cited at appropriate places. All 
photographs are original, and published here for the first time. Col- 
leagues V.N. Orlov and LP. Mitina of the Department of Vertebrate 
Zoology at Moscow State University provided great assistance during 
the preparation of this book. 

We here once more acknowledge with thanks all the above-men- 
tioned persons, organizations and others who helped in the production 
of this book. The authors especially acknowledge Prof V.A. Popov 
(Kazan') and Prof N.I. Larina (Saratov) who provided great help in 
the preparation and reading of the manuscript, and who made many 
useful suggestions. 

All work was to a great extent accomplished in the Zoological 
Museum, Moscow State University, whose rich collections were used. 
The authors express their gratitude to prior and present directors of 
this Museum: Professors S.G. Soin and N.A. Gladkov. 

The first volume "Mammals of the Soviet Union" was received in 
our country, and in other countries as well, with great appreciation 
and was officially and unofficially reviewed and put out in German 
translation. The authors hope that this volume will also be acceptable, 
even though they are quite sure that their work is not free from spe- 
cific and general errors. The authors kindly request those who use 
their book to inform them of any errors noticed and to send necessary 
corrections to the following: Professor Vladimir Georgevich Heptner, 
Zoological Museum, Moscow State University, Herzen St., No. 6, 
Moscow K-9 [U.S.S.R.]. 



Table of Contents* 



FOREWORD TO THE ENGLISH EDITION v 

FOREWORD ix 

SYSTEMATICS OF THE CLASS MAMMALIA xix 

KEY FOR IDENTIFICATION OF ORDERS xxi 

ORDER SIRENIA ILLIGER, 1811 1 

Suborder Trichechiformes Hay, 1923 18 

Family Hydrodamalidae Palmer, 1895 18 

Genus Hydrodamalis Retzius, 1794** 18 

Hydrodamalis gigas (Zimmermann, 1780) 18 

(Steller Sea Cow) 

Description and Biology 19 

Geographic Distribution and History of Extinction 41 

ORDER CARNIVORA BOWDICH, 1821 51 

Key for Identification of Families in the Order Camivora 64 

Family CANIDAE Gray, 1821*** (Wolves and allies) 66 

Key for Species Identification of the Wolf Family 74 

Subfamily Caninae Gill, 1872 78 

Genus Nyctereutes Temminck, 1839 (Raccoon Dog) 78 

Nyctereutes procyonoides (Gray, 1843)**** 82 

Diagnosis 82 

Description 83 

Systematic Position • 87 

Geographic Distribution 87 

*Pages 1000-1001 in Russian original. The English table of contents is not a 
literal translation. 

**Palmer, 1895 in Russian original — Sci. Ed. 

***Fisher, 1817 is now considered to have priority — Sci. Ed. 

****1843 in Russian original — Sci. Ed. 



XVI 



Geographic Variation 91 

Biology 92 

Practical Significance 120 

Genus Canis Linnaeus, 1758 (Wolves) 124 

Canis aureus Linnaeus, 1758 (Jackal) 129 

Diagnosis 1 29 

Description 1 29 

Systematic Position 132 

Geographic Distribution 134 

Geographic Variation 140 

Biology 141 

Practical Significance 160 

Canis lupus Linnaeus, 1758 (Gray Wolf) 164 

Diagnosis 165 

Description 1 66 

Systematic Position 175 

Geographic Distribution 175 

Geographic Variation 182 

Biology 198 

Practical Significance 262 

Genus Alopex Каир, 1829 (Arctic Fox) 271 

Alopex lagopus (Liimeaus, 1758) (Arctic Fox) 274 

Diagnosis 275 

Description 275 

Systematic Position 279 

Geographic Distribution 279 

Geographic Variation 287 

Biology 292 

Practical Significance 376 

Genus Vulpes Oken, 1816* (Foxes) 385 

Vulpes сапа Blanford, 1877 (Afghan Fox) 390 

Diagnosis 390 

Description 390 

Geographic Distribution 393 

Geographic Variation 394 

Biology 395 

Practical Significance 395 



*Frisch, 1775 is now considered to have priority — Sci. Ed. 



XVI 1 



Vulpes corsac (Linnaeus, 1768) (Corsac Fox) 396 

Diagnosis 397 

Description 397 

Geographic Distribution 402 

Geographic Variation 406 

Biology 409 

Practical Significance 465 

Vulpes vulpes (Linnaeus, 1758) (Red Fox) 469 

Diagnosis 471 

Description 472 

Systematic Position 480 

Geographic Distribution 482 

Geographic Variation 485 

Biology 504 

Practical Significance 561 

Subfamily Simocyoninae Zittel, 1893* 566 

Genus Cuon Hodgson, 1838 (Red Wolf) 566 

Cuon alpinus (Pallas, 1811) (Red Wolf) 571 

Diagnosis 571 

Description 571 

Systematic Position 574 

Geographic Distribution 574 

Geographic Variation 578 

Biology 580 

Practical Significance 586 

Family Ursidae Gray, 1825** 586 

Key for Species Identification of the Bear Family 596 

Genus Ursus Linnaeus, 1758 (Bears) 597 

Ursus (Ursus) arctos Lirmaeus, 1758 (Brown Bear) 601 

Diagnosis 604 

Description 604 

Systematic Position 612 

Geographic Distribution 613 

Geographic Variation 632 

Biology 649 

Practical Significance 676 



*Dawkins, 1868 is now considered to have priority — Sci. Ed. 
**Fischer, 1817 is now considered to have priority — Sci. Ed. 



XVIU 



Ursus (Thalarctos) maritimus Phipps, 1774* 

(Polar Bear) 678 

Diagnosis 678 

Description 679 

Systematic Position 684 

Geographic Distribution 685 

Geographic Variation 694 

Biology 700 

Practical Significance 712 
Ursus (Selenarctos) thibetanus G. Cuvier, 1823 

(White-chested, Black Bear) 713 

Diagnosis 713 

Description 713 

Systematic Position 719 

Geographic Distribution 720 

Geographic Variation 723 

Biology 724 

Practical Significance 733 

t LITERATURE CITED 

t INDEX OF LATIN NAMES OF TAXA** 



*1774 in Russian original — Sci. Ed. 

**The Russian original also included an Index of Russian Names of Taxa, omitted 
here — Sci Ed. 

t These Sections appear at the end of part lb. — Gen. Ed. 



Systematics of the Class Mammalia 



9 The classification of major subdivisions (orders) adopted in the present 
publication is based on the systematic monograph of the class by 
Simpson (1945), who relied extensively on paleontological data. It is 
very close to the system of classification proposed by M. Weber (1928), 
which in its time was well founded on the anatomy of extant forms. 
The orders proposed by Simpson have long been widely recognized 
and their grouping into higher taxa raises no serious objections. 

Extreme division of the orders is a recent trend. Thus, marsupials 
have been divided into three orders — polyprotodontids-Polyproto- 
dontidia, caenolestids — Caenolestia, and diprotodontids-Diprotodontia, 
cetaceans into two (toothless whales — Mystacoceti and toothed 
whales — Odontoceti), primates into three (lemurs — ^Prosimiae, sim- 
ians — Simiae, and tarsiers — ^Tarsioidea), and so on. All these proposals, 
however, are not yet well founded. Moreover, among mammalian 
systematists extreme division is not generally favored. Neither is it 
commended here. An order should be primarily an integrating and not 
a divisive concept. Otherwise the same situation will arise in mam- 
malogy as has already occurred in ornithology, where the concept of 
an order already has largely lost its significance; as a result the sci- 
entific aspect of systematics of vertebrates has suffered greatly. 

In the classification of orders under consideration here, only one 
change has been made; i.e., pinnipeds have been treated as an inde- 
pendent order and not as a suborder of Camivora. Based on the 
systematics of recent mammals, these two groups separate naturally — 
they are not less isolated than some other orders. 

Based on already established views, Lagomoфha are treated here 
as an independent order (Weber considers them only a suborder of 
rodents). Hence, instead of the 18 orders of Simpson, we propose 19. 
Of these, 10 (52.6%) are represented in our fauna, one — the sea cow — 
is now extinct (Steller's sea cow). 

The main subdivisions within the orders in most cases adhere to 
Simpson's scheme. By and large his classification is the most well 



XX 

founded. Nevertheless, there are a series of deviations, some signifi- 
cant, mainly in inteфretation of the content of the genus, which is 
understood more broadly than is generally the case. In our opinion 
this conforms better to the structure and principles of natural classi- 
fication. 

The systematic classification adopted is given in the chart below. 
Orders present in the USSR fauna are marked with an asterisk. (V.H.) 



10 



CLASS MAMMALIA 



Subclass PROTOTHERIA 


Order 




MONOTREMATA 


Subclass THERIA 




Infraclass METATHERIA 


Order MARSUPIALIA 


Infraclass EUTHERIA 






* Order INSECTIVORA 




Order DERMOPTERA 




* Order CHIROPTERA 


Cohort UNGUICULATA 


Order PRIMATES 




Order EDENTATA 




Order PHOLIDOTA 



Cohort GLIRES 
Cohort MUTICA 



* Order LAGOMORPHA 

* Order RODENTIA 

* Order CETACEA 



rSuperorder FERAeF* Order CARNIVORA 
|_* Order PINNIPEDIA 



Cohort 
FERUNGULATA 



Superorder 
PROTUNGULATA 



Superorder 
PAENUNGULATA 

Superorder 
MESAXONIA 
Superorder 
PARAXONIA 



Order 
TUBULIDENTATA 



Order PROBOSCIDEA 
Order HYRACOIDEA 

* Order SIRENIA 

* Order 
PERISSODACTYLA 

* Order 
ARTIODACTYLA 



11 



Key for Identification of Orders 



1 (2). Hind limbs absent; fishlike body with large bilobate caudal 

fins set horizontally CETACEA. 

2 (1). Hind limbs present; body not fishlike and tail, if present, 

not in the form of a bilobate fin. 

3 (4). Forelimbs in the form of leathery wings 

CHIROPTERA. 

4 (3). Forelimbs of a different structure. 

5 (6). Fore- and hind limbs very short and resemble paddles, 

i.e., all the digits entirely enclosed in a common 
integument PINNIPEDIA. 

6 (5). Fore- and hind limbs of a different structure, not paddle- 

shaped. 
7 (10). Feet possess hooves. 

8 (9). Only one hoof on each limb PERISSODACTYLA. 

9 (8). On each limb, two large hooves and two small ones above 

them ARTIODACTYLA.' 

10 (7). Hooves absent (claws present). 

11 (14). Between large chisel-shaped incisors and molars on jaws, 

a large toothless gap (diastema) occurs; its length not less 
than length of entire row of molars on corresponding jaw. 
Canines absent. 

12 (13). Two incisors on upper jaw RODENTIA. 

13 (12). Four incisors on upper jaw; behind each large shaф incisor 

is located a minute blunt tooth LAGOMORPHA. 

14 (11). Diastema between incisors and molars absent or much 

smaller than length of molar row. Canines present. 



' Limbs differ in structure in camels. The leg terminates in a broadened callused 
pad and true hooves are absent. There are two greatly broadened nails. Camels have 
not been included in this table as they are domesticated animals (wild camels are 
extinct).* 

*lt is now believed that wild populations of Bactrian camels still survive in China 
and Mongolia — Sci. Ed. 



XXll 



15 (16). Anterior portion of muzzle extended into well-developed 

small proboscis. Anteriormost tooth on each jaw, or only 
upper jaw, much larger than adjacent tooth.^ FNSECTIVORA. 

16 (15). Anterior portion of muzzle does not form a proboscis. 

Anteriormost tooth on each jaw not larger than one next to 
it CARNIVORA.3 



'If the structure of teeth is different, proboscis is always present and furthermore 
the front leg extremely short, feet very broad with huge claws, and set on edge with 
inner surface facing backward (moles). 

Юп the coast of the Commander Islands, skulls of the extinct Stteller's cow — 
a member of the order of sea cows (Sirenia) — have been found. They are distinguished 
by upper and lower jaws measuring 60 cm in length, in which teeth or traces of them 
(alveoli) are absent (fig. 6). 



,3 ORDER OF SEA COWS, 

OR SIRENIANS 

Ordo Sirenia lUiger, 1811 



Superorder of Probosciforms^ 

Superordo PAENUNGULATA Simpson, 

1945 

Order of Sea Cows, or Sirenians 
Ordo SIIIENIA Illiger, 1 8 1 1 * * 



15 Mammals of large size, entirely monomoфhic,*** highly special- 

ized as typical aquatic animals. The species of this order represent one 
of the extreme degrees of specialization among mammals. 

Body elongated, fusiform, and in general appearance, more similar 
to that of large dolphins, but as compared to the latter, and in relation 
to total animal size, the head is smaller and located on a short, movable 
neck not noticeable externally in all animals. In connection with the 
low mobility of the animal, the general body constitution is much 
heavier than that of dolphins and the body diameter in its thickest part 
is much larger. There are no external hind limbs. The hind part of the 
body is elongated and thins gradually, rather than abruptly, into a 
relatively thick tail region which turns into a wide, flattened horizon- 
tally oriented tail fluke, lacking intemal skeleton (other than the vertebral 
column and the connective tissue). The posterior edge of the fluke 
forms a convex arch and the whole fluke is blade-like (Family 
Trichechidae — manatees), or else the posterior edge is deeply incised 
in the middle and the tail is bilobed (remaining families). 

The anterior extremities are in the form of true flippers, without 
any [external] signs of differentiation into digits. Some species have 
nail rudiments on the II-V fingers (several manatees). The flippers are 
freely mobile in the shoulder joint and (in contrast to cetaceans) pos- 
sess a considerable mobility in the elbow and hand articulations (see 
p. 30). The animal leans upon them, raises the anterior part of its body, 

*In Russian original, slonoobraznykh; literally, "of elephant-like form" — ^Sci. Ed. 
**Co-edited by Daryl Domning (D.D.), Howard University, Washington, D.C. 
***Except Dugong — -D.D. 



moves, and performs other relatively complicated movements (arrang- 
ing food in its mouth, etc.). 

On the head, in connection with the peculiar structure of the skull 
(see below), the upper lip is extremely developed, forming with the 
nasal region a massive fleshy mass hanging down over the lower, 
normally developed lip, and represents a sort of wide, blunt and short 
proboscis. The mouth opening lies on the lower surface of the head. 
The shape of the "proboscis" varies in different forms (higher or wider), 
but in all forms it is highly movable, capable of changing its shape, and 
plays an important role in collecting food. In some animals (for ex- 
ample the manatees, Trichechus), the actively movable lateral parts 
can even mimic to a degree, a prehensile organ. It acts like the index 
finger and thumb of humans, and by bending it to the left and to the 
right the animal can grasp plants. The nostrils are located on the an- 
terior upper part of the "proboscis"; they are crescent-shaped and closed 
(open only during inhalation and exhalation). The eyes are small and 
round; eyelids are devoid of eyelashes but richly supplied with special 
glands, secreting abundant viscous fluid that protects the eyes from the 
influence of water. ' The Harderian gland is highly developed but, in 
contrast to cetaceans, the lachrimal glands are apparently absent. A 
sclera is not formed.* A well developed and highly movable nictitating 
membrane is present. The ears are located far behind the eyes and are 
represented by very small orifices; a pinna is completely absent. 

The hair cover of the body is completely, or almost completely, 
absent and is represented by fine hairs scattered singly on the body, and 
having the structure and, apparently, function of sensory organs. This 
fulfills the need for high tactile sensibility of the skin, important for an 
animal living among aquatic vegetation in coastal zones. In the fetus, 
the number of these hairs is considerably greater. On the upper lip and 
partway around the mouth are found a large number of hairs; usually, 
however, they are short and do not form the "whiskers" characteristic 
of some pinnipeds. The majority of the hairs are elastic and bristle-like, 
but some are softer. Some hairs have a tactile function while others, 
especially the coarser ones, are considered a device for assisting the 
grasping of food with the upper lip. 

'In animals removed from the water, this fluid accumulates in the corner of the 
eye and then flows out. The legend of the "weeping sirens" has originated from this, 
and so does the name lamantin (from the French Lamenter: to weep). This oily liquid 
is, apparently, a product of Meibomian and Harderian glands. 

*Meaning not clear — Sci. Ed. 



In the majority of species, the skin has a very thick connective 
tissue layer (about 20-25 mm) and very thin epidermal layer (cuticle 
is about 0. 1 of the total thickness of the skin). The skin [in manatees] 
is thicker on the sides than on the back as is characteristic of mammals 
in general; on the abdomen, it is approximately half as thin as that on 
the sides, but on the back, it is thicker than on the abdomen. Sometimes 
(Steller's sea cow, see below), the epidermal layer is especially devel- 
oped and strong. In some species (Amazon manatee, Trichechus 
inunguis) the border of the caudal fin bears a hard comification border 
(or is included in it) which may be detached and again re-formed.^ The 
fatty layer is well developed. 

The [skin] color is dark, more or less evenly brown, brownish or 
black in color; from below, it is lighter in some animals, and sometimes 
of a rare-meat color. 

There are two mammary glands. 

The female genital opening is slit-like, placed fairly far in front of 
the anal opening. The testicles lie in the abdominal cavity (far anterior, 
near the diaphragm); the male copulatory organ is long with highly 
developed софога cavernosa, lies hidden under the skin, and contains 
no bony structures. 

The skulls of all species of this order are completely monomor- 
phic,* very peculiar and differ greatly in general structure from those 
of other mammals. They have some similarities with the skull of artio- 
dactyls and perissodactyls, and particularly with those of elephants and 
hyraxes, but they shaфly differ, in general and in details, from the 
skulls of cetaceans. 

In its general features, the skull is characterized by the fact that the 
nasal openings are greatly shifted and directed upwards, and the pre- 
maxillae are large in size. In some animals they are very large and 
massive and are bent downwards in their anterior half (symphysial 
part), taking on a beak-like form, and giving the skull a curved form 
17 in side view (Hydrodamalidae, Steller's sea cow; and especially 
Dugongidae, dugong). In other forms, the premaxillae are less massive 
and extend forward, but are not, or are only slightly bent downwards 
(Trichechidae), and do not fiise together in a symphysis.** The nasal 
processes of the premaxillae extend far backwards, reaching the nasal 
bones. Together with the enlarged maxillary bones, they delimit the 

-Judging by some figures of Steller's sea cow (see below), they apparently 
possessed such a structure on the posterior border of the fin. 
* Except Dugong — D.D. 
**Not true; symphysis is formed — D.D. 



nasal cavity. The nasal bones are very small and shifted far posterior, 
forming [part of] the rear wall of the wide nasal opening, on the bottom 
of which the widely opened vomer is found. The nasal bones are some- 
times absolutely rudimentary [or absent]. The maxillae are large, and 
the palate is long. 

The supraoccipital is large, greatly extending forward, but not 
wedged between the parietals. The suture between the parietals disap- 
pears at an early stage and a single parietal is formed, constituting the 
cranial roof The frontals are separated and with large supraorbital 
processes. The squamosal bone has a large zygomatic process, to which 
the jugal bone is connected. The space between the exoccipital and 
squamosal is only partially filled by the petrosal bone (petrosum); [the 
remaining space] consists of a large opening (filled with connective 
tissue in the living animal) leading to the cranial cavity. The tympanic 
bone (tympanicum) is quite closely connected [fused] to the petrosal 
bone, forming the petrotympanic bone (petrotympanicum). The tym- 
panic bone is in the form of a fairly massive half-circle, but it does not 
form a bony auditory capsule (bulla). The petrotympanic bone is not 
fused with the skull but freely articulates and connects with the squa- 
mosal bone. The anterior and posterior lacerate foramina (for. lacera) 
are fused in a common large one. The lachrymal bone is small, not 
perforated, and in some animals it is connected with the surrounding 
bones. There is no lachrymal canal. The palate is long and the palatine 
bones, being small and shifted posterior, play a very insignificant role 
in its composition. 

The mandibular bones are massive and heavy, connected in an 
elongate symphysial portion; they form a single bone. The anterior part 
of the lower jaw is recurved downwards, and the degree of its bending 
corresponds to the form and position of the premaxillae (most strongly 
in the dugong, and to a somewhat lesser degree in Steller's sea cow). 
The ascending ramus of the lower jaw has strong, well developed 
articular and coronoid processes, directed forward and of various rela- 
tive size in various forms, and a massive broad angular process. There 
are no air cavities in the skull. 

The dental system is highly peculiar and differs greatly from that 
of other mammals. The cheek teeth [in Trichechus]* are absolutely 
identical with a short crown carrying two transverse crests, each of 
which is formed from three tuberosities. During occlusional wear, the 
masticatory surface is flattened. The number of simultaneously func- 
tioning teeth in one jaw ramus of a mature animal is 3—^ or even 2 

*D.D. 



(Dugong) and from 5—6 to 8 (manatees, Trichechus). All the teeth are 
considered as true molars [some are deciduous premolars]\ and inci- 
sors, premolars and canines are absent. Only in male [and a few old 
female]^ dugong one permanent incisor erupts on each side of the upper 
jaw, converted into a peculiar small (up to 10 cm) "tusk", directed 
downward. 

With wear, the teeth [in TrichechusY are replaced by new ones, but 
not by the old tooth being pushed out by the new one growing under it. 
The newly formed teeth appear at the extreme rear of the toothrows and 
at the same time the anterior ones ("old" teeth) fall out, the crowns having 
almost wom out and lost their roots, which are well developed in "younger" 
teeth. Accordingly, the toothrow is continuously shifted forward. This 
1 8 takes place because the posterior side of the bony septum between teeth 
(i.e. the septum between each two successive teeth) is resorbed, while 
on its anterior surface (i.e. adhering to the rear surface of the tooth in 
front) this bony septum grows forward. In manatees, replacement of 
teeth occurs throughout the whole life and the number of molars is, as 
a matter of fact, not limited. In manatees, for example, in every jaw 
quadrant ("half jaw) not less than 30 teeth or possibly more^ are 
replaced throughout the animal's life. If we take into consideration only 
the teeth functioning at a given moment, the dental formula for a fully 

mature dugong (Dugong) is I— C — P — M— =10, while in the case of 
ь ьу & &j 000 2 

110 3 3 
young animals of this species, it is DI— I— C— DP— M— = 28*. 

The biological significance of this unusual method of tooth replace- 
ment in manatees probably lies in the fact that in feeding on aquatic 
vegetation much sand** is obtained and the teeth are wom out rela- 
tively quickly. 

'D.D. 

'This manner of tooth replacement from behind forwards occurs only in 
elephants.*** However, the number of teeth replaced in the course of life does not 
exceed 6 and only one tooth is functioning in each jaw ramus (in all 4 cheek teeth). 
Sirenians and elephants are also similar in that the tusks in both are formed by the 
incisors. The transverse orientation of the crests on the teeth in sirenians is similar 
to the corresponding structure in primitive Proboscidea. 

*In Russian original, "I— C— P— M— = 26" which is erroneous — ^D.D. 
10 3 3 

**Probably siliceous phytoliths in the vegetation — D.D. 
***And some macropod marsupials — D.D. 



Overall, a reduction in the dental system occurs in this order, and 
in one species (Steller's sea cow, Hydrodamalis) teeth are lost com- 
pletely. 

This reduction is connected with, and probably due to, the fact that 
in all species, in both the upper and lower jaws there are large elon- 
gated homy plates occluding with each other in front of the tooth rows. 
On the lower jaw, the plate is located on the symphysial part which is 
bent downward, while on the upper jaw, the plate lies on the lower 
concave surface of its anterior part. Such a system is used for seizing 
and grinding food and has no analogue among other mammals. The 
small comified part lying in the region of the anterior end of the 
premaxilla in even-toed ungulates has, however, some ftinctional and 
moфhological similarities to the upper plate in sirenians (see vol. I). 

Owing to the great reduction of the skeleton of the posterior ex- 
tremities, true sacral vertebrae are absent,* and not differentiated frorn 
the lumbar vertebrae; the caudal region is long. In the genus Trichechus, 
the number of cervical vertebrae is six.'* The number of trunk vertebrae 
in various species is 19—24; caudal — 22—29, and 35 (Hydrodamalis, 
Steller's sea cow); ribs, 14-19 pairs.** Thoracic vertebrae are very 

*True of manatees, but not dugongs — D.D. 

"The number of the cervical vertebrae in other species cannot apparently be 
considered established. According to Weber (1928) there are 7, while Mohr (1957) 
stated that in the dugong and Steller's sea cow there are "?6 ?7". Steller (1745— 
1753), who gave a detailed description of the Steller's sea cow anatomy, noted 6 
cervical vertebrae. 

The skeleton in the Zoological Museum of Moscow University has six vertebrae 
(V.G. Heptner), but it is possible that one is missing. In the skeleton found in the 
Zoological Museum of the Academy of Science (Leningrad) are seven cervical vertebrae 
(K.K. Chapskii), but it cannot be excluded that long ago in assembling the skeleton, 
some vertebrae might have been added to reach the number normal for mammals. A 
skeleton present in the Museum of Kiev University has seven vertebrae, but it is possibly 
more correct to consider them six; one thoracic vertebra may be placed with the cervical 
ones (A.P. Korneev). It is unlikely, however, that Steller could commit such a terrible 
mistake, and therefore it subsequently was agreed to consider the number of the cervical 
vertebrae in this animal as being six until further special investigation is carried out. 

A solution of this problem based on museum skeletons is at the present time 
difficult, because all of them are actually composed of bones from different individu- 
als. Only Evermann (1893) bought in 1891 an almost complete skeleton from the 
Commander [Islands] which was found "a short time" before that. The finder con- 
firmed that the skeleton was found not far from the water's edge under a few inches 
of sand. The skeleton contained the complete set of cervical vertebrae which, after its 
study in the National Museum of Washington, was shown to be 7. 

The deviation in the number of cervical vertebrae is a rare exception among 
mammals. Besides manatees, only the two-toed sloth, Choloepus hoffmanni, has 6 
vertebrae, and the three-toed sloth, of the genus Bradypus, has 9 vertebrae. 

**14— 19 pairs among living species — D.D. 



few and only 2-3 or 5 of the anterior ribs (Steller's sea cow) are 
connected to them, while the remaining ribs are free. Clavicles are 
absent. 

The scapula is well developed and carries a normal spine. The 
articulation of humerus with scapula is quite free, the elbow joint is 
freely movable and the wrist joint has a considerable degree of mobil- 
ity. Ulna and radius are separate and only in a few forms do they fuse 
distally in older animals.* In some young animals the whole set of 
сафа1 bones is present, but in mature ones, they fuse together into two 
or three large bones. The manus is pentadactyl and of normal construc- 
tion; Polydactyly is not found, and supernumerary phalanges do not 
normally occur in the species. Little is known, in contrast to cetaceans, 
of the structure of the forelimb skeleton in limiting their capability for 
different movements, such as acute bending of the hand in conveying 
food to the mouth, etc. 

The skeleton of the hind limbs is utterly lost and only in a few 
species is there a totally insignificant rudiment of femur with an articu- 
lating head. The pelvis is represented by only the ischium and ilium, 
which are united together [in dugongids] into a rod-shaped bone in 
which its constituent elements are not well differentiated. There is no 
acetabulum. 

The bones of the skeleton have no marrow cavities, and all of them, 
including those of the skull, are characterized by what is termed 
pachyosteosclerosis — ^Haversian canals are closed, all bone tissue is 
very compact resembling that of elephant bones, and shows conchoidal 
fracture. Moreover, the bones are relatively massive. All these features 
make the skeleton very heavy. Apparently, these features are of adap- 
tive significance for animals living in coastal waters, bringing their 
specific gravity close to that of water, thus facilitating underwater 
submergence, etc.^ 

The kidneys are lobular as in true marine animals (whales, pinni- 
peds), the intestine is very long and exceeds the body length by 13—20 
times, the large intestine is very long, only a bit shorter than the small 
intestine, and sometimes longer (Steller's sea cow). The caecum is well 
developed and in some species has accessory diverticula. The stomach 
is a complex organ, with a shaф constriction between the cardiac and 
pyloric parts, an additional blind sac or outgrowth in the cardiac region, 
and two blind sacs in the pyloric part.** 

*Not true; in most they fuse both proximally and distally — D.D. 
^Paleontological data show that pachyostosis appears in the evolutionary 
sequence during the transition to life in water. 
**Actually part of duodenum — D.D. 



10 

The uterus is bicomuate and the placenta zonal. 

With the exception of some differences in size and development of 
tusks in males [and females] of one species (Dugong), there is no 
sexual dimoфhism. 

The body length of smaller species reaches 200—300 and may 
reach 360 cm. Body diameter, in its thickest part, is about 150—155 cm 
(for a length of 203—247 cm) and the weight is up to 400-600 kg. In 
the largest species (Steller's sea cow), the body length reaches 800 cm, 
the diameter of the trunk about 620 cm and the weight up to 3200 (200 
poods) and 4000 kg (moфhological structure according to Weber, 1927, 
1928; Mohr, 1957 and other references, and material in the ZMMU 
[Zoological Museum, Moscow University]). 

They are specialized aquatic animals which do not come out on 
land,^ and are associated with shallows of coastal rivers, in particular 
with calm bays, lagoons, and river mouths, and with rivers rich in 
aquatic vegetation. They do not live far from shores, and particularly 
[do not inhabit] pelagic parts of the sea. Three species are purely 
marine, one marine and freshwater,* and one is strictly freshwater. 
Although most are inhabitants of tropical, warm water, one species 
inhabits the cold temperate zone. They are herbivorous forms, feeding 
20 on submerged and floating aquatic plants ("grazing") along the shores. 
They are very sluggish, flaccid, and clumsy animals [on land] of quite 
low mental level. They are usually found in small groups. Young are 
usually single. 

This order is widely distributed, the major part of its range being 
characteristic of littoral (coastal) marine animals and only in part char- 
acteristic of the ranges of animals associated mainly with the lower 
courses of larger rivers. Therefore, everywhere the range is typically 
linear (Heptner, 1936). The general range is divided into four isolated 
parts, differing greatly in their size — two lie in the Atlantic and two in 
Indian and Pacific oceans. These parts are not continuous, but divided 
into smaller and larger sections. 

In America, the range (reconstructed, including transgressions in 
the north) extends to the southern coast of North Carolina (Wilmington)** 

''If a sea cow is exposed on land or subjected to drying in some other way (for 
example, ebb-tide), it does not die of this, but as do whales, is "crushed" by its own 
weight and is overheated.*** However, these animals cannot move themselves on 
land. 

*Two considered marine, and two both marine and freshwater — D.D. 

**To New England — D.D. 

***No evidence for this — D.D. 



12 

and occupies the coasts of the Gulf of Mexico, islands of the West 
Indies, and Central America, as well as part of the northern and north- 
eastern coast of South America — the shores and river mouths in 
Colombia, Venezuela and Guiana to Cape Norte in the east (slightly 
southeast of Maraca Island, Brazil), lying somewhat north of the mouth 
of the Amazon. The range of this [partly] marine species of the West 
Indies {Trichechus manatus) does not, apparently, reach the mouth of 
this river.^ Moreover, the range in South America (of the freshwater 
species T. inunguis) covers the basin of Orinoco* and Amazon basin 
far to the west including, in any event, the region of a right-hand 
tributary of the Amazon, the Rio Madeira (according to the data of 
Hatt, 1934; G. Allen, 1942; Cabrera, 1960).^ 

At the present time the range is strongly modified and animals have 
been exterminated in most places, and north along the Atlantic coast, 
manatees are no longer found beyond the southern extremes of Florida,* 
and where they are protected. 

In West Africa, the range occupies coastal lagoons and river mouths 
from the Senegal River on the north to the Cuanza River in northern 
Angola. The species inhabiting this region {T. senegalensis) lives in 
fresh [and salt] water and they penetrate upstream in the majority of 
West African rivers, especially the large ones. Thus, in the Niger, their 
range includes its left tributary, the Benue, but in the Niger itself, the 
range extends almost to Timbuktu. Manatees are widely distributed in 
the Congo basin.** Their upstream distribution along rivers is generally 
restricted by waterfalls and rapids. Reports and suggestions concerning 
the occurrence of manatees in Lake Chad and its basin are mistakes 



''The range suggested by Cockrum (1962) is, apparently, not true even for the past. 
There are direct indications that owing to hydrologicai conditions (Muфhy, 1936 and 
others), manatees did not occur to the east beyond the place mentioned, and were not 
found on the coast of northern Brazil east of the mouth of Amazon River. They did not 
reach Cape Sao Roque (the eastern point of the continent) and have not spread to the 
south, although, in the area southward from the Cape, along the eastern coast, the 
conditions for them were apparently favorable.*** Cockrum (1962) assumed that the 
area of distribution of these animals extended along the coast of South America to the 
mouth of the La Plata [River]. 

"^''Rio Madevia" and "Rio Madeva" given by Mohr (1957) are probably misprints 
instead of "Rio Madeira" (type locality for this species). 

♦Not true — D.D. 

**No evidence for this statement — D.D. 

***West Indian manatees occur in this region today — D.D. 



13 

lacking foundation,* as are data about the presence of manatees in the 
Great [Rift] Lakes of Africa.^ 

In the Indian and southwestern parts of the Pacific Ocean, the range 
of the dugong {Dugong dugon) occupies the greatly extended shoreline 
22 from East Africa and the Red Sea in the west to eastern Australia, the 
Solomon and Marshall islands, and the Ryukyu islands. The range in the 
above-mentioned areas was probably discontinuous from time immemo- 
rial, but because of inadequate information it is now impossible to 
reconstruct the true range. The range is divided at present into separate 
parts. '° Along the eastern African coast, the range extends in the south 
to Delagoa Bay at 26° S. Lat., a little south of Lourengo Marques, but 
in earlier times it may have been extended to the south (to the Cape of 
Good Hope?), and occupied the whole Red Sea along the southern coast 
of the Arabian peninsula, though not east of Aden. It occupies the coast 
of Madagascar, mainly the west and the small islands off the African 
coast (Mafia** and others)." 

Yet, the dugong is apparently absent and probably has not been 
present for a long time in the majority of places in the north of the 
Indian Ocean; i.e. the southern coast of the Arabian peninsula, in the 
Persian, Oman, and Bengal gulfs, and the shores of Indostan.*** The 
range here (the second part of the Indo-Pacific Ocean range) occupies 
only the most southerly part of the west coast of India, starting from 
Cananora (Canara [Cannanore], about 12° N. Lat.) southward and to 
the southernmost part of the eastern coast of the peninsula lying adja- 
cent to Ceylon, which is included in the range. 

The third part of the Indo-Pacific Ocean region occupied by sea 
cows is connected with extreme southeastern Asia, Australia and the 
islands lying between them. In the south, the range includes the western 
coast of Australia to the north of Shark Bay (about 25° S. Lat., a little 

*There is now some evidence that manatees occur in the Lake Chad basin — D.D. 

''Information concerning the occurrence of manatees in the past (about 1800) at 
Saint Helena Island in the Atlantic Ocean, which in its time provided material for 
zoogeographical speculation, is false and is probably based on the appearance of 
elephant seals (Hatt, 1934; МифЬу, 1936). 

'"The range described here differs considerably from that presented by Cockrum 
(1962, map) which was incorrect at least in part. For example, the occurrence on the 
coast of Madagascar was not given although a series of reports were available. 

**Off coast of Tanzania — Sci. Ed. 

"Concerning the Mascarene, Comoro and other islands of the Indian Ocean and the 
Mozambique Strait associated with Madagascar, there are no reports. Only Cockrum 
(1962) includes the Mascarene islands in the range. 

*** Archaic; Indian subcontinent — Sci. Ed. 



14 

south of Carnarvon), the northern coast, and the eastern coast south to 
Botany Bay (near Sydney). In the west, the range covers the Greater 
and Lesser Sunda islands. In the east, it covers the Solomon and Marshall 
islands (there are no definite records from other islands of Micronesia 
or Melanesia, in particular the Bismarck archipelago). In the north, the 
range includes Sumatra, the coasts of Malacca Strait (no definite records 
are known from the Nicobar and Andaman islands), the coast of the 
Gulf of Siam, the whole of the Philippine archipelago as far north as the 
Batan islands, Taiwan and the Ryukyu archipelago north to Amami-o- 
shima (Amamioshima, 28° N. Lat.). Thus the range includes the southem 
coast and western extremes of New Guinea (concerning the northern 
coast, records are lacking), and the Molucca Island group. In this re- 
gion, the range occupies the Coral, Arafiira, Timor, Banda, Java, Flores, 
and Celebes seas. Macassar Strait, part of the South China Sea, i.e. all 
the internal seas lying between Asia and Australia, part of the East 
China Sea, the extreme eastern coast of the Indian Ocean, the extreme 
west part of the Pacific Ocean, and that part in the Micronesian region 
(Blanford, 1876; Allen, 1942; Carter, Hill, Tate,* 1946; Cockrum, 1962). 

A distinct, very small part of the range lies separated from other 
parts in the Commander Islands [Hydrodamalis only]. 

A known event is the passage of a dugong through the Suez Canal 
to the Palestinian coast (Aharoni, 1930). 

Remains of Tertiary sirenians (fam. Dugongidae) are known far 
from the present limits of the range — in Africa, Europe and even in 
Jamaica. 

Sea cows, as was previously mentioned, are considered as a shaфIy 
characterized group, and thus, accepted by all modem investigators as 
an order. However, a short time ago, the sea cows were sometimes 
23 considered as a suborder (Sirenia) of the Order Subungulata, which, 
besides Sirenia, includes Hyracoidea, Proboscidea and the extinct 
Embrithopoda. Systematic and phylogenetic connections between all 
three contemporary groups is unquestioned — ^as was reflected in their 
combination in the Superorder Paenungulata (see p. xx;. However, dif- 
ferences between them, even by evaluating all palaeontological data, are 
those normally recognized for ordinal rank. The differences between 
Sirenia and both Proboscidea and Hyracoidea are undoubtedly greater 
than those between Pinnipedia and Camivora. 

The systematic position of this order is, according to recent data, 
determined by their previously mentioned relation to Paenungulata. This 

*Misspelled "Teg" in Russian original — Sci. Ed. 



15 

order has, apparently, more common features with the order of elephants 
and they can be considered as sister groups although the relation be- 
tween them is quite distant. The nature of tooth replacement and some 
of its peculiarities, which is not elsewhere found among mammals, the 
transformation of incisors into tusks, some features of skull appearance 
and others, are among these common characters.* The similarity some- 
what increases in the comparison of the more primitive ancient forms of 
both groups. Sea cows are sometimes considered closer to Hyracoidea. 
In any case, there is no doubt (and this is very obvious) that the sea 
cows have no relation either to pinnipeds or to whales. All the features 
common with whales are no more than convergences, and close study 
reveals deep principal differences between them. The greatest similarity 
with whales lies only in the general appearance, the absence of the rear 
extremities, and in some details characteristic of aquatic mammals in 
general. 

In the eighteenth century, at the beginning of the last century and 
somewhat later, even competent authors (Pallas, for example) included 
the sea cows in the order Cetacea (whales) as a special group ("grass- 
eating whales"). However, even at that time, there were some indications 
not only of the independence of sea cows as an order, but also of their 
relatively closer relation to elephants. This point of view was, a long 
time ago, generally accepted, though it must be emphasized that the 
systematic isolation of this order is very shaф and that its relation to 
elephants is very distant. The same also applies to ungulates, though in 
the system of orders of the class, sea cows are always put between 
elephants (or hyraxes) and one of the orders of ungulates [see p. xx, this 
volume]. 

The order comprises two suborders: Trichechiformes, in which to- 
gether with the extinct ones, all living forms are included, and 
Desmostyliformes [now an order], including only extinct forms (one 
family with perhaps five genera). The systematic position of this group 
is, however, not completely clear, and some taxonomists consider it a 
separate order, Desmostylia, which is close to Sirenia. 

The suborder Trichechiformes comprises five families, of which two 
(Prorastomidae — one genus, Protosirenidae — one genus) are extinct and 
three living: manatees, Trichechidae; dugongs, Dugongidae and kapustniks, 
or Steller's sea cows, Hydrodamalidae.** The family Trichechidae is, 



*In part, no longer considered true — D.D. 

**Now reduced to a subfamily of Dugongidae — D.D. 



16 

apparently, considered the least specialized among the existing ones. Spe- 
cies of this family have a relatively less developed, only slightly depressed 
anterior ("premaxilla") part of skull [rostrum], complete dental system, and 
6 cervical vertebrae. The tail is blade-like and single-lobed. This group 
includes one genus, Trichechus. The family Dugongidae comprises four 
subfamilies: Halitheriinae (7 genera), Miosireninae (1 genus) and 
Rytiodontinae (one genus) are extinct, and the Recent Dugonginae with 1 
genus Dugong. This last family is characterized by the considerable reduc- 
tion of cheek teeth (to 2), the presence of tusks in the upper jaw of males, 
the extremely enlarged and shaфly depressed upper jaw, and 7 cervical 
vertebrae. The tail is bilobed. The family Hydrodamalidae, kapustniks or 
Steller's sea cows, with 1 [recently extinct] genus, Hydrodamalis, is char- 
24 acterized by the complete loss of teeth, a fairly highly developed and 
depressed rostrum, and 6* cervical vertebrae. The tail is bilobed.'^ 

Evidence on the origin of the group is given by representatives of 
the above-mentioned extinct families existing only in the Eocene, and 
the family Dugongidae which is represented by its extinct subfamilies 
in the Eocene — beginning especially from the Middle Eocene to Plio- 
cene. The other existing families have no extinct representatives, or only 
their Pleistocene remains are known (manatees). 

The oldest Middle Eocene forms, such as Eotheroides, though show- 
ing some essential differences from the existing ones (complete dental 
system, apparently, since they were characterized by the absence of 
homy plates, a fairly well developed pelvis, etc.), were true sea cows. 
They are well linked with the existing forms by means of a succession 
of forms. In the Middle and Upper Eocene, they were already very 
widely distributed and, therefore, the origin of this order must be related 
to the Paleocene. As was shown, the similarity in the structure of skull 
and teeth of the more ancient sea cows with primitive proboscideans and 
hyraxes is more evident than in the existing sea cows. The ancestors of 

*Now known to be 7 — D.D. 

'^The above-mentioned classification of the order differs from that adopted by 
Simpson (1945), mainly in the acceptance of Hydrodamalidae as a family (which he 
considers as only a subfamily of Dugongidae). This is partially determined by the fact 
that Steller's cows possess 6 cervical vertebrae and not 7.* Other authors consider this 
group also as a family (Weber, 1928; Mohr, 1957) even when there was the assumption 
of 7 vertebrae. The order of the families as regards increasing specialization is also 
changed. In sum of characters, the family Hydrodamalidae is closer to Dugongidae than 
to Trichechidae and is considered as a fiirther development of this type. The difference 
in the number of cervical vertebrae shaфly separates the families from one another. 

A more thorough analysis of the extinct and existing forms and their relationship 
is given by Simpson (1932). 



17 

sea cows must, probably, be searched for among land forms, especially 
those which gave rise to the other Paenungulata, i.e. among the hyraxes 
(order Hyracoidea) and proboscideans (Proboscidea) as well as ungu- 
lates (Artiodactyla and Perissodactyla). However, direct "basal forms" 
of the order are not known. 

The existing species in the order are 5. In the genus Trichechus L., 
1758 (= Manatus Brtinn.) there are 3 species: T. manatus L. (shores of 
the Americas — see above), T. inunguis Natt. (basins of Amazon and 
Orinoco*) and T. senegalensis Linlc (sea coasts and the lower reaches 
of the rivers of West Africa). In the genus Diigong Lac, 1799 
(= Halicore 111.) there is 1 species — D. dugon Erxl.** (the coastal 
region of the Indian Ocean and the adjacent southwestern Pacific). The 
kapustniks, or Steller's sea cows, Hydrodamalis Retzius, 1794 — 1 
species — H. gigas Zimm. (coastal waters of the Commander Islands). 

The greater number of species are endemic to the Atlantic Ocean, 
with two associated with its western coast and one with its eastern 
coast. One species is endemic to the Indian Ocean and small parts of 
the Pacific contiguous to it. The practical value of the group is insig- 
nificant. They are only hunted locally for their meat. In many places 
these animals have been extiфated and their number is limited almost 
everywhere. One species was completely exterminated (Steller's sea 
cow). 

The USSR fauna comprises only 1 species, the kapustnik or Steller's 
sea cow, Hydrodamalis gigas Zimm., 1780 — ^which constitutes 20% of 
the species of the order and about 0.3% of the number of species of our 
fauna (V.H.). 



*Not found in Orinoco — D.D. 
**Now attributed to Mullet — D.D. 



Suborder of True Sirenians 

Subordo TRICHECHIFORMES Hay, 

1923 

Family of Kapustniks, or Steller's sea cow'' 
Familia HYDRODAMALIDAE Palmer, 1895 

Genus of Kapustniks, or Steller's sea cow 
Genus Hydrodamalis Retzius, 1794 



25 1794. Hydrodamalis. Retzius. Kongl. Svenska Vet.-Acad. Nya Handl. 

Stockholm, 15, p. 292. Hydrodamalis stelleri. 
181 1. Rytina.'^ Illiger. Abhandl. d. Berlin. Akad., 1811. p. 64. Rytina 

borealis. 
1814. Nepus. Fischer. Zoognosia, 8, p. 640. Nepus stelleri. 
1822. Stellerus. Desmarest. Mammalogie, pt. 2, p. 510. Stellerus 

borealis. (V.H.). 

KAPUSTNIK, SEA COW, STELLER'S COW'^ 

Hydrodamalis* gigas Zimmermann, 1780 

1780. Manati gigas. Zimmermann. Geograph. Gesch d. Menschen und 

d. vierf. Thiere 2, p. 426. 
1785. Manati balaenurus. Boddaert. Elenchus anim., p. 173. 



'^The characteristics of the family and genus are not given here, as they are clear 
from the characteristics of the order and the species description given below. Only the 
main generic synonyms are given. 

'"•The first (and correct) spelling of the genus name. Rhylina is a later emendation. 

''The name "kapustnik" [from cabbage] and "sea cow" are vernacular. The first 
originates from "sea cabbage" — a. name which was and is still given to some seaweeds 
used as animal food. 

"Steller's cow" is a literary name translated from a name of German origin. 

*Misspelled Hidrodamalis in Russian original — Sci. Ed. 



19 

1788. Trichechus manatus, var. (3. borealis. Gmelin, Syst. Nat., 1, p. 

61. 
1794. Hydrodamalis stelleri. Retzius, Kongl. Svenska. Vet.-Akad. Nya 

Handl. Stockholm, 15, p. 202. 
1815. Rytind cetacea Illiger. Abhandl. d. Berlin. Akad., 181 1, p. 70.'*' 

(V.H.). 

Description and Biology''' 

26 The body length of an adult animal exceeds 720 cm. The dimensions of 
the female killed on Bering Island on June 12, 1742 were as follows'^: 

"Body length from top of upper lip to end of 

right lobe of caudal fin 741.5 cm 

Distance from top of upper lip to mouth angle 39.5 cm 

Distance from top of upper lip to shoulder 132.5 cm 
Distance from top of upper lip to genital opening 490.0 cm 

Length of genital opening 26.0 cm 
Tail length from constrictor muscle of anal 

opening to beginning of lobes of caudal fin 192.5 cm 

Head circumference at level of nostrils 78.0 cm 

Head circumference at level of eyes 122.0 cm 

Neck circumference at occiput 204.0 cm 

'^The Commander [Bering] Island is the type locality of all of these names, based 
on the description of Steller. 

'The following description of the body structure and habits of the animal, written 
between quotation marks and without reference to author, is a translation from Steller 
(1751, 1753), done for this book.* This text is accurate and almost complete. Only a 
few parts, of interest only for Steller's contemporaries (concerning identity of his species 
to the American one, the possible movement of the animal on land, its use in 
medicine . . . etc.), the greater part of the anatomical section, some measurements (Steller 
gave 42) and other redundancies were left out. Data of other authors are given with 
reference to their sources. 

The complete work of Steller, written in Latin and almost simultaneously published 
in German, has not yet appeared in Russian. Only small parts of it were recast or 
published in the form of free translations by S. Krasheninnikov (1755) who had at his 
disposal Steller's manuscript. Steller's text was not always consistent. He wrote his 
famous work in Kamchatka or on the return journey to Petersburg. Steller died along 
the road through Siberia in Tyumen in 1746. It is possible, judging by the accuracy 
with which he kept his diary (1793), that at least part of the work was written on the 
island (V.H.). 

"^Translated from Paris feet (V.H.). 

*But see also Jordan, D.S., 1899— D.D. 



20 



Height of anterior part (proboscis) 
Trunk circumference at shoulder 
Maximum trunk circumference halfway 

along posterior part of the abdomen 
Tail circumference at point of divergence of 

lobes of caudal fm 
Distance between ends of lobes or width of tail 

across lobes 
Height (thickness) of tail lobe 
Length of entire internal (upper) lip area which is 

covered by vibrissae or lip broom 
Its width [lip broom] 
Width of external upper lip in chin direction with 

its declined surface which is entirely covered 

with long white bristles 
Its height [external upper lip] 
Width of the lower lip, which is dark, without hairs 
|or bristles, smooth, drooping on breast, 

heart-shaped 
Its height [lower lip] 
Distance from lower lip to chest 
Measurement or width of mouth from one end 

to other 
Width, or rather length, of stomach 
Length of intestine from mouth to anus 
Accordingly it is 20.5 times longer than the 

animal itself (body length is more than twenty- 
four English feet as shown from the first 

measurement). 
Distance from genital opening to constrictor muscle 

of anal opening 20.4 cm 

Diameter of trachea below larynx 10.7 cm 

Length of heart 56.0 cm 

Width of heart 64.0 cm 

Length of kidney 80.0 cm 

Width of kidney 45.6 cm 

Length of tongue 30.6 cm 

Width of tongue 9.0 cm 

Length of teats 10.2 cm" 



21.0 


cm 


367.0 


cm 


622.0 


cm 


143.0 


cm 


199.0 


cm 


26.5 


cm 


13.2 


cm 


7.7 


cm 


35.7 


cm 


25.5 


cm 


19.7 


cm 


17.3 


cm 


138.0 


cm 


52.0 


cm 


112.0 


cm 


1521.0 


cm 



21 







*l^.. » . ,i. -— » ■— Ч 



27 Fig. 2. Depiction of the icapustnik (Steller's seacow) Hyrf/Wawa//5g/ga5 Zimm., 
from the chart of the first officer of V. Bering's "Saint Peter", Lieutenant 
S. Waxell, in 1742. The drawing was probably made by G. Steller or is a 
copy of his drawing (from Pekarskii, 1869).* 

27 According to Steller the number of vertebrae is 6 cervical, 19 

thoracic, and 35 others. The skeleton present in the Zoological Museum 
of Moscow University has 6 cervical, 19 thoracic, and 32 lumbar, sacral 
and caudal vertebrae. 

The skull dimensions of aduh Steller' s sea cows according to the 
material of the Zoological Institute of the Academy of Sciences and the 
Zoological Museum of Moscow University, are as follows: condylobasal 
length (12 specimens) 638-M 673-722 mm, zygomatic width (17 speci- 
mens including measurements of 8 specimens published by Stejneger, 
1883-1884) 324-M 344-373 mm. The condylobasal length of the lower 
jaw (14 specimens) 432-M 465-491 mm. 

Males were, apparently, somewhat larger than females and accord- 
ing to Stejneger (1883-1884) possessed somewhat wider zygomatic 
arches.** Thus his material (8 specimens) was divided into two groups: 
in one of them (probably males) the zygomatic width was larger than 
half.the skull length from the upper end of the foramen magnum to the 
anterior point of the premaxillae, and in the second (females), these 
measurements are the same. 

The length of the masticating plate (specimen in the Zoological 
Institute of the Academy of Sciences) is 182.4 mm, its width anteriorly 
is 80.5 mm, and posteriorly 81.5 mm. Its color is dirty, dark brown (in 
living animals the masticating plate was white in color — see below) 
(V.H.). 

*Probably drawn by Plenisner; see Stejneger, 1936 — D.D. 
**No specimens are of known sex — D.D. 



22 

". . . The famous Clusius* considered this animal (Steller thought 
that he worked with the American "manati", V.H.) clumsy and very 
ugly, because he saw only the hide stuffed with straw. The living animal 
was, actually, sufficiently distinctive. However, on account of its exter- 
nal appearance, and the style of movement, this animal is extremely 
striking. It has so thick a skin that it resembles the bark of an old oak 
tree rather than animal skin. The skin is black, rough and wrinkled as 
if covered with small pebbles, or resembles a shagreen, and is hard, 
elastic and devoid of hair; an axe or Ьафооп can hardly pass through 
it. Its thickness is about an inch (25.5 mm, V.H.) and, in transverse 
section and because of its color and smoothness, it greatly resembles 
ebony (black, V.H.) wood. This external cover is, however, not the true 
skin but rather its external layer (epidermis). On the spine the skin is 
smooth or bare. In contrast to this, it is covered, from the occiput to the 
tail lobes with annular wrinkles, making the skin somewhat uneven. The 
sides are very rough, as if covered with pebbles, carrying many con- 
vexities looking like sponges without peduncles (?). The skin looks 
disgusting, especially on the head. The described upper layer of the skin 
invests the whole body like a shell. Its thickness frequently reaches an 
inch and consists entirely of tubules, as in Spanish rattan or in Hindu 
and Chinese bamboo. These tubules adhere closely to each other as if 
united, and are so peфendicularly located that they can be separated 
along their whole length. Each fiber ("hair", V.H.) lies and is fixed in 
28 the dermis at its lower end, is somewhat rounded, swollen and thick- 
ened, and has a tuberiform bulb, and therefore, the cut-off pieces of this 
upper skin have [when seen] from below many swellings like Spanish 
skin (Morocco ? V.H.). On the contrary, the upper surface of the dermis 
lying under it is, as in a thimble, entirely covered with small depressions 
in which the bulbs of the epidermis lie. Since these so-called tubules lie 
very close to each other, they are always wet and swollen, but if this 
upper layer of the skin is horizontally cut off, nothing is excreted from 
the tubules and the cut-off surface is entirely smooth, similar to a cut 
piece of claw. However, when these pieces are dried in the sun, cracks 
are formed and the whole piece may be broken like a piece of bark and 
the whole structure of the skin, in the form of distinctive tubules, is quite 
obvious. Through these tubules, a liquid or watery mucus is excreted, 
which is more on the head and side and less on the back. If the animal 
lies for some hours on the dry shore the back becomes completely dry, 

*C. Clusius, 1605, Exoticorum libri decern, Leiden — D.D. 



23 




Fig. 3. Depiction of Steller's sea cow from "Zoographia", P.C. Pallas (1811). 

The drawing is, probably, a copy from the figure of S. Waxell (see Fig. 2) and 

described by Pallas as a "draft". 



29 



while the head and side remain wet or moistened. Thus, the thick upper 
layer of the skin on the animal performs two functions: when the animal 
occupies places with shaф stones, or when in winter it is always found 
in the midst of the ice, it does not strip off its dermis during feeding, 
or when strong waves strike the animal against rocks and submerged 
stones, which I myself often saw, it does not die, but on the contrary, 
is protected as with armor. On the other hand, in summer and as a 
consequence of the very high evaporation, the animal does not lose its 
heat and in winter, with extreme cold, it can survive. This occurs be- 
cause the animal cannot stay under water for a long time, like other 
water animals and fishes, but during its feeding, half its body must be 
above water, and subjected to the effect of cold. 

According to my observation of many dead animals that were cast 
up by the sea, these animals died because their external skin layer was 
damaged by rocks; this occurs mainly due to the ice in winter. 

In addition to this, I have frequently noticed that when the wounded 
animals were pulled by haфoons to the bank, they struggled very furi- 
ously by their abdomen and tail while being supported by their forelimbs, 
and thus large pieces of the external skin layer were detached. I also 
noticed the detachment of these layers from the forelimbs or hands and 
from the so-called hoof (hand, V.H.) as well as from the lobes of the 
tail, and this confirmed my opinion . . ," 

"... some sea cows had on this upper skin layer fairly large white 
spots and stripes and therefore the skin appeared speckled and this color 
penetrated through to the true skin [dermis]. 

The upper skin layer on the head, around the eyes and ears, on the 
mammary glands and arm-pits, possessed a tuberculous (stony) form or 
the form of shagreen, and was thoroughly filled with harmful insects 



24 




Fig. 4. Depiction of Steller's sea cow assembled by J.F. Brandt (1846) and 

identified by him as an "imagined picture" ("Icon idealis"). The skeleton of the 

animal was not known at that time and Brandt had at his disposal only fragments 

of the skull and masticating plates. 

which grow round the skin.^^ Very frequently, the latter appeared to be 
perforated by these insects, and even the true skin [dermis] was injured. 
If this occurs, the oozing lymph of the penetrated glands, in which there 
are usually vesicles containing fat, forms large, thick protuberances 
similar to those on whales, and such patches make the body ugly. 

Under this previously described upper layer of skin there lies around 
the whole body the true (actual) skin [dermis]. Its thickness is two lines 
(about 5 mm), soft, of white color, very compact and firm, resembling 
that of a whale in respect to its structure and thickness, and perhaps 
required for the same puфoses. 

The head, as compared to the huge body, is small, short and where 
it ends is not evident; it is long and almost quadrangular in form; 
however from the crown to the lower jaw, it broadens. The crown itself 
is flattened, and its black superficial skin has bumps (stony, V.H.), or 
looks like shagreen and appears almost broken; it is thinner (by 1/3) 
than in other places, and is easily separated. From the occiput to the 
nostril, the surface is bent and again bent from the nose towards the 
nostril as far as the lips. The anterior proboscis (the protruding part of 
the upper lip, V.H.) has a length of 8 inches (20.5 cm) and, from the 
nose to the occiput, it is strongly thickened. 

The shape of the mouth (opened mouth) does not pass backwards, 
but is located along the sides. The external lip is so large, and declines 
to the angles of the face and extends far above the lower jaw, that if 
one examines only the head, he thinks that the shape of the mouth 
opening extends backwards, but at a very low level. 



"An amphipod crustacean — Cyamus rhytinae Brandt. (V.H.). 



25 

The opened mouth is small in relation to the animal's size; how- 
ever, it need not be large because the animal feeds only on seaweeds 
and lives on them. 

Both upper and lower lips are paired, and divided into external and 
internal regions. 

The external upper lip protrudes obliquely forward as the most 
anterior part of the proboscis and has a semicircular form. It is flat- 
tened, swollen, thick, 14 inches (36 cm) wide and 10 inches (25.5 cm) 
high, white in color, smooth and covered with a large number of tu- 
bercles or cones. From the middle of each tubercle grow white transparent 
bristles, 4-5 inches (10—12.5 cm) long. 

The internal lip is 5 inches (14 cm) long, 1/3 inch (8 mm) thick,* 
completely separated from the external one and united with the external 
lip by its base only; it hangs over the palate like a calf s tongue (!), 
is all tattered and is hard, like dry twigs of a broom. The above- 
described upper lip tightly closes the mouth from above, is mobile, and 
30 is used to browse on seaweeds in much the same way as horses and 
bulls collect their food by sliding the lips apart and extending them 
forward. 

The lower lip is also paired. Externally it is black, smooth, without 
bristles, of irregular heart-shaped form, and sometimes looks like a 
chin, seven inches (18 cm) wide and six and eight-tenths inches (17.5 
cm) in height. Internally, the lower lip is only slightly separated from 
the external one, is rough and when the mouth is closed, it is not seen 
at all because the arch-shaped external lip lies under it; however, the 
internal lower lip adheres to the internal upper one and tightly closes 
the mouth. 

Where the lower jaw adjoins the upper one, the space remaining 
between them is occupied by numerous dense and thick white bristles 
one and one-half inches (38 mm) long. This prevents anything from 
being dropped from the mouth during mastication, or washed off by sea 
water: the water, which together with food is always taken into the 
animal's mouth, is drained outside when the mouth is closed. 

The above-described bristles have the thickness of a goose feather 
quill, white, hollow, and with follicular swellings from below. By them, 
it is clear to see without a magnifying glass how nature forms our hairs. 

When the animal lies directly on its abdomen, the height of the 
anterior part of the proboscis or muzzle, measured peфendicularly 



Trom Steller (1753)? Steller (1751) reads "2y inches wide"— D.D. 



26 

from the nostrils to the lips, is equal to eight inches (20 cm); it extends 
from the nose to both the anterior-most part of the lips, and to the 
lateral side of the upper jaw, like mesenteries (in folds), anteriorly 
rounded, then thickens and its size greatly increases. The external lips 
are very greatly enlarged, thick, as if swollen and, as in cats, carry 
many white pores (!) or orifices. From each, thick white bristles protrude 
which become thicker as they gradually approach the mouth opening. 

The thickest bristles are those growing between the lips of both 
jaws. They use these bristles in browsing seaweeds, and they prevent 
anything from falling from the mouth. The lower jaw is shorter than the 
upper one and is movable. The lips of both upper and lower jaws are 
mobile and function as in our draft animal [horse]. When the animal 
picks vegetation with its hands or forefeet from the plants growing 
underwater on the stony bottom, it cleans the fronds from the hard stems 
and roots (which are not utilized as food) with the above-described 
bristles as if cut off with a blunt knife. These stems and roots are thrown 
up by the waves during [high] tide and heaped on the shore in very large 
piles, indicating that an animal is present. Since the stems of aquatic 
plants are much harder and thicker than those of terrestrial plants, these 
beasts must have harder, more compact lips than any terrestrial animal. 
Therefore, such lips are not utilized by man as food and they are not 
softened by boiling or any other means. The inner lips are so con- 
structed that, if they are cut open, separate small areas are seen on them 
similar to those on a checkers or chess board. These are innumerable, 
very small and thick red muscles of rhomboidal or trapezoidal shape. 
Between these muscles numerous areas resembling tendons are located, 
and form a net or tissue of vesicles filled with liquid fat. These lips 
easily release all their fat when cooked in boiling water and when they 
are defattened, all the white areas look like the numerous tendons of a 
net. This natural structure, it seems to me, has three puфoses, namely: 

1 ) in order to make the lips stronger and more compact and protect 
them from external injury; 
31 2) in these muscles the heads and tails, i.e. their beginnings and 

ends [origin and insertion] are so located that their heads are slopingly 
directed toward the mouth opening while their tails, or ends, are located 
obliquely in relation to the parietal, in such a way that their beginnings 
and ends form a sort of garland and this helps raising and moving the 
heavy lips; 

3) with the assistance of this structure, the lips can be, to a certain 
degree, bent in a helix or spiral and since the whole head is covered by 



27 



a thick cortex it bends with difficulty and thus, when the animal wishes 
to browse on elastic seaweed, it is not necessary to move the whole 
body. 

Mastication in these animals occurs in a manner different from 
other animals. It is achieved not by teeth, because they possess no teeth, 
but with the assistance of 2 long bones found in their place. These bones 
[masticating plates] are large, of white color and are considered as 
whole rows of teeth or as one tooth mass. One of these bones is fixed 
to the palate, and the other to the lower jaw. 

These bones are, however, attached in an entirely unusual manner 
and the method of their attachment cannot be attributed to any of the 
known ones. They cannot be described as fixed because they do not 
insert in the jaw as do teeth, but their protuberances and pits lie in 
corresponding protuberances and pits in the palate and lower jaw. 






Fig. 5. Depiction of Steller's sea cow made from the so-called "Royal Agriculture 
Manuscript" of S. Waxell and his later (compared to that described by Pekarskii, 
1869 — see Fig. 2) chart (Bikhner [Biichner], 1891). In the original, the upper 
drawing has a length of 26.5 cm (in that of Bikhner, 1891 — from the nose to the 
engraved tail 20 cm), and two times more than the lower drawing. The upper 
drawing ("Royal Agriculture Manuscript No. 2") is, apparently, a more detailed 
drawing of the same animal in Fig. 2. The lower drawing ("Royal Agriculture 
Manuscript No. 1 ") was greatly changed during re-drawing, probably by the same 
source (according to Bikhner, 1891 — this is a depiction of another animal). 



28 

Moreover, the dental bones are fixed in the anterior part of the hard skin 
of the inner upper Ир, along the sides of the mouth in the edge of the 
bones, and are finally inserted into the palate and lower jaw in the 
posterior part by means of a double outgrowth, and that is how they are 
attached. 

These grinding bones occupy the place of all molar teeth and, like 
a thimble or sponge, have many small holes on the lower surface through 
which, as in the teeth of [other] animals, the arteries and small nerves 
32 pass dorsally; these dental bones are far from smooth and covered with 
numerous bent wavy grooves; between them prominences are raised 
which, in their turn, occlude during mastication, into the grooves of the 
opposite dental bone and between them, and the marine plants are 
ground as in a hand or fulling mill. 

The nose lies on the uppermost part of the head and to the sides as 
in a horse; the two nostrils are separated by the nasal septum of one and 
one-half inches (38 mm) in width. The nostrils themselves are 2 inches 
(5 1 mm) long and the same in diameter; they are open, but inside, there 
are many flexures or blind pockets. The inner walls of the nasal open- 
ings are extraordinarily thick, wrinkled and covered with compact skin 
with many black pores which have bristles protruding from each of 
them. These bristles have the thickness of sewing thread and are one- 
half inch (13 mm) long and are easily detached; they perform the function 
of those in other animals. 

The eyes lie exactly mid-way between the end of the muzzle and 
the ears, at the same height as the upper part of the nose or slightly 
higher. They are relatively small in comparison to the large body of the 
animal, and are not larger than those of sheep. Externally, they have no 
eye-lashes or any similar structure surrounding the eyes, and they pro- 
trude from the skin through a rounded hole, which according to my 
measurements, has a diameter of scarcely one-half inch (13 mm). The 
iris is black in color and the eyeball is yellow blue. The comers of the 
eyes are not noticeable; moreover, when the skin around the eyes is cut 
off, a cartilaginous body like a cock's comb protrudes at the large 
(anterior) comer, as in the case of the sea otter. When needed, this body 
closes the eye, nearly in the same way as in animals which are grazing, 
i.e. searching in dusty and dirty soil, and has such a membrane (third 
eyelid, V.H.), which can be stretched by the animal on the eyes to 
prevent the entrance of any harmful objects. This cartilaginous body 
forms, with its other side, the wall of the lachrimal sac and is connected 
with it by a common compact skin. On cutting into the lachrimal sac. 



w 



i 




z 



I 



^fciP-TfflB 



29 

dense mucus is found inside; the lachrimal sac is large enough to allow 
the entrance of a chestnut, and it is lined by glandular skin. 

Externally, the ears are represented by a small, open orifice but 
there are no traces of external ears. The ears themselves are unnotice- 
able, unless a person searches carefully for them. They cannot be 
differentiated from the surrounding skin which resembles shagreen. The 
shaft of a feather can hardly pass into the ear. 

The internal ear canal is smooth and coated with an extraordinarily 
smooth (polished) black skin, and therefore it can be easily detected, 
when the muscles of the posterior part of the head are separated, by its 
color. 

The tongue is 12 inches in length (30.5 cm), its width three and a 
half inches (9 cm), and as in cows, it is pointed at its end; its surface 
is rough as a file and covered with low warts. The tongue is so deeply 
placed in the mouth that one may doubt its presence. If it is pulled by 
hand from the mouth, its anterior part fails to reach the grinding plate 
of the mouth by one and one-half inch (38 mm). However, if the tongue 
were as long as in other animals, it would interfere with the above- 
mentioned wide bones during mastication. 

The head is not visibly demarcated from the neck, and these are 
similarly connected to the trunk without any demarcation in any place, 
and thus the body is constructed as in a fish. The neck is not clearly 
demarcated, except that it is half the length of the head, elongated, 
round, and somewhat more slender than the posterior part of the head. 
In spite of that, the neck is movable due to the presence of movable 
vertebrae. However, these movements are only observed during feeding 
in living animals. During this, the animal lowers its head as does an ox, 
grazing in a dry pasture. However, if the animal lies resting or after 
being killed, the animal seems to have an immovable neck owing to its 
33 thick clumsy outer skin and especially due to the absence of any exter- 
nal trace of the cervical vertebrae. 

From the scapula to the umbilicus, the trunk is again strongly thick- 
ened; and from the latter to the anus, the trunk size decreases and it 
again becomes slender. The flanks are as bulging as the belly itself, and 
the latter, due to the great weight of the intestine included in it, is very 
swollen, stretched and distended by the intestine. From the umbilicus to 
the anus, the trunk is thinned to the same degree as from the chest 
(mammary glands) to the neck. 

In spring and autumn, when the sea cow is in fat condition, the back 
is somewhat convex, and in winter when they are emaciated, the back 



30 

appears flat, and on both sides of the vertebral column there appear deep 
hollows; and thus the vertebral column and vertebrae are clearly seen. 

The ribs are curved up on both sides of the spine as a convex arch 
and in those places where they articulate with the vertebrae, as in the 
case of human beings, they again bend down forming an arch, where 
on both sides of the vertebral column they form a double groove along 
the whole length of the back. 

The tail starts at the 26th vertebra; the caudal vertebrae are 35. 
Posteriorly from the anus and in the direction of the tail fins the tail 
becomes thinner and not completely flat, but rather somewhat quadran- 
gular ... the tail muscles fill the empty places in the vertebrae and the 
angles between their processes by 4 plates and therefore, the tail looks 
like a long quadrangle with blunt angles. 

The tail is generally thick, extremely strong, very hard, stiff (slightly 
movable) and ends in a black fluke. The latter is not divided into rays 
but forms a single unit. Its material is similar to the whalebone (baleen) 
which is used by the dressmakers, and, similarly, is composed of leaves 
lying one above the other as rings of a tree or the grain of a piece of 
wood. At a distance of one-quarter from the end, it appears wider and 
somewhat resembles a coarse spindle*. . . . The fluke itself, forming 
the tail end, has a width or length of 78 inches (199 cm), a height of 
7 3/10 inches (18.5 cm), and a thickness of 1.5 inches (38 mm). . . . 

The caudal fluke somewhat resembles pincers (?), with horns or 
hands of equal size. . . . The tail blades are located, as in whales and 
dolphins, to the sides, and are not, as in the majority of fishes, in a line 
with the spine. If the sea cow slowly moves its tail to one side, it slowly 
swims forward, but if it strongly moves the tail upward and downward, 
it rapidly swims forward and tries in this way to move away from 
harmful hands wanting to pull it on to the shore. 

The most remarkable thing in which this animal differs from all 
terrestrial animals which live in water. and from aquatic animals which 
spend some time on land, and from all animals in general, is its entirely 
unique hand or fore limb, if it can be so called. At the neck, two limbs 
originate straight from the shoulder bones, 26 inches (68 cm) in length 
and consisting of two parts. The uppermost bone [humerus] is con- 
nected to the scapula by a flat articulation. 

Here, as in the case of humans, there are radius and ulna, both of 
which border on the carpal bones with the metacaфal bones (called 

*ln Russian original, "osti kolosa"; literally, grass awn — Sci. Ed. 



31 

metatarsals and premetatarsals, V.H.), but neither fingers, nails nor 
claws can be detected. The above-mentioned anterior and middle (meta- 
tarsal and premetatarsal) parts of the limb are covered with dense fat 
and coated with many ligaments and superficial skin, and have the form 
of the newly overgrowing skin on a wound on the human extremity. 
However, the skin and especially its superficial layer is thicker on that 
part and also harder and drier, so that the ends of the hands roughly 
resemble a kind of crustacean claw [nail] or horse-hoof, even though 
the horse-hoof is зЬафег and narrower and therefore more adapted to 
34 digging, or scraping. At the posterior border, these limb ends are smooth 
and curved; from below, they are, to a certain degree, concave (empty) 
and covered with numerous dense bristles about one-half inch (12—13 
mm) in length and, when scratching, are as shaф as the twigs of a 
broom. In one of these animals I noticed nipper-like limbs, divided into 
two parts, similar to the cow hoof. However, such division was not an 
actual one and was restricted to the superficial skin layer, and was 
probably accidentally brought about and not naturally created in this 
form. This is still more possible, as the superficial skin layer lying on 
the so-called claw handle is very dry and may be easily cracked. 

This Platonic man, as the famous Raiyus (Ray, V.H.)* said in jest, 
used its hands in different manners. It can use them as fins in swim- 
ming. It walks on them in shallow waters as on legs; it holds itself upon 
them, resting among the slippery submerged stones and rocks, and 
scrapes and tears off the sea plants and seaweeds from the rocky 
bottom as do horses with their forelimbs. It can lean upon them and 
thus make great resistance when wounded by a haфoon and dragged 
from water to the shore. In such conditions, it holds so strongly that the 
upper layer of skin on the hand is split and torn off in pieces. During 
sexual intercourse, the female lies on her back and swims in this 
position. The male lies upon her; she holds the male strongly and he, 
in his turn, embraces her. 

On the anterior part of the breast, two teats are found, located in 
a manner different from that in other animals. Each one lies under the 
corresponding arm as in the case of humans and has a similar appear- 
ance. Each breast has a length of 1 1/2 feet (45 cm), is convex, and 
bears many folds or spiral flexures directed inwards. The breast is 
glandular, extremely hard, harder than the cow udder and without fat 
deposits. The fat layer covering the whole body is found also on the 

*John Ray— D.D. 



32 





Fig. 6. Skull of Steiler's sea cow. Upper figure, side view of skull; lower, 
dorsal view (from Stejneger, 1884).* 

breast and is as thick there as on other parts. However, the upper layer 
of skin is thinner, softer and more wrinkled. The nipple also is of dark 
outer skin, with circular wrinkles. However, this superficial skin layer 
is soft; the nipple hangs under each hand or shoulder; in lactating 
animals, it has a length of 4 inches (10 cm) and a thickness of 1.5 
inches (38 mm). If the lactation period is over or if the animal has not 
yet given birth, the nipples do not protrude and are so short that they 
can be considered as accidentally occurring warts. At that time, the 
breast also is not very swollen. The milk of these sea cows is as sweet 
35 and fatty as regards its taste and density, as that of sheep. Frequently, 
I drew milk from nipples after the death of the animals, as from cows. 
The area around the nipple is very wrinkled and a little raised above 
the surface of the breast. In each nipple there are from 10 to 12 milk 
ducts which open into the teat. The boiled mammary glands were 
somewhat harder than a cow's udder and they had a weak gamy taste. 

* Actually from Simpson (1932) — D.D. 



33 

These animals mate in the same way as human beings. The male 
lies above and the female lies below. The penis is 32 inches (81.5 cm) 
in length; it is tightly fixed to the belly wall by its sheath, reaching the 
umbilicus; it is thick and ugly in its external appearance. It is similar 
to that of the horse and has the same head, but is somewhat larger. 

The female genital opening is located eight inches (20 cm) in front 
of the anus. Anteriorly, the opening is nearly triangular ... to the rear 
it narrows. . . . The anus ... is closed by a constrictor muscle, not very 
tightly. Its width is four inches (10 cm); the sphincter is white. . . ." 

From the biological observations of Steller given in the part of his 
work entitled "Description of behavior and nature of this animal", the 
following lines deserve particular attention (many interesting observa- 
tions are not directly related to the biology of this beast). "... I was 
lucky enough, or rather unfortunate, to be obliged to observe the be- 
havior and customs of this animal in front of my door for ten months^° 
and thus I wish briefly to state that I can explain [their behavior] with 
trustworthiness. 

These animals like shallow and sandy places on the shore. How- 
ever, they very willingly stay where rivers and creeks pour into the sea, 
and on their banks, as they are attracted to the fresh water coming to 
the sea; therefore, they stay there in groups. When they feed, they lead 
their young or the newly bom ones in front of them, following them 
from behind and on the sides, and very carefiilly surround them, keep- 
ing them in the center of the herd. When the tide rises, they come very 
near to shore, so that I could not only beat and prick them, but some- 
times even stroke them with my hand on their back. If they suffer from 

36 a severe pain, they only moved away further from the coast than usual, 
but they soon forget that and come near again. ^' 

37 Usually, their whole families are found close to each other; the 
male with the female ("with his wife"), with one adult [offspring?] and 
also some other young. Each husband, apparently, does not have more 

38 than one wife; they give birth to young at any time of the year, but most 
often, judging by the newly bom young, in autumn. Proceeding from 

-"Bering's ship "Saint Peter" escaped a rock and was very lucky to be thrown on 
the flat shore of a broad bay, where the crew spent the winter in mud huts and where 
Bering himself died. This place was very favorable habitat for sea cows (V.H.). 

^'Steller writes elsewhere ". . . it is rather possible to believe that it is possible to 
make this animal tame, rather than this strange cunning which is attributed to it in 
stories (spoken about the American species, which Steller considered the kapustnik, 
V.H.). Due to its extraordinary stupidity and greediness for food, this animal is tame by 
nature, and there is no necessity to domesticate it . . .". 



34 



'm 




35 Fig. 7. Bering Island, shore near Cape Manati. Former habitat of Steller's sea cow. 

Photograph by S.V. Marakov. 

the fact that I observed their mating early in spring, I think that preg- 
nancy lasts more than one year, and that they give birth to no more than 
one young, which can be deduced from the short uterus and the fact that 
the adult female has only one pair of nipples. Moreover, I have never 
observed more than one calf beside its mother. 

These insatiable animals eat continuously. They are extremely greedy 
for food, so they hold their heads all the time under water; accordingly, 
they very rarely worry about their life and safety, and it is easy in a 
boat or even swimming naked among them to choose which one is 
needed to remove from the sea. During feeding they do nothing but 
thrust their nose out from water every four or five minutes, and, making 
a noise similar to the neighing, sniffing, and snorting of horses, they 
breathe out air with a small amount of water. During feeding, they put 
one leg after other and in this way they swim slowly, or walk like oxen 
or sheep. Half of the trunk, mainly the back and flanks, are always 
protruding from the water. During feeding, sea-gulls sit on their backs 
and pick the lice (crustaceans, V.H.) found on the upper layer of the 
skin and in this v/ay they are useful to them, in the same way as crows 
do on swine and sheep. The sea cows do not eat all the types of sea 
weeds without selection, but they mainly eat: 1) wrinkled cabbage, like 
Savoy cabbage with latticed leaves; 2) water plants with upper parts 
similar to a cudgel; 3) plants which look like the whips of ancient 



35 

Romans; 4) very long sea weeds with a wavy vein edge.^^ In places 
where these animals were feeding, even for one day, the roots and 
stems thrown out by the sea lie on the shore in large heaps. After they 
feed to satiety^^ some of them lie on their back. To avoid being on dry 
land during ebb-tide, they swim away from shore farther into the sea.^'* 
In winter, these animals frequently die under the ice which floats along 
the coast, and dead animals are cast up onto the shore by the sea. The 
same also happens when the waves strongly strike the rocks, taking 
these animals with them. In winter the animals are so emaciated that 
all their ribs can be counted along the vertebrae. 

In spring, they come together as do human beings, mainly in the 
evening when the sea is calm. However, before copulation they show 
many amatory signs in their relation to one another. The female slowly 
swims in the sea here and there, and the male follows her all the time; 
the female moves around for a long time near the male until she reaches 
extreme sexual desire. She then lies on her back as if she is tired and 
does that seemingly unwillingly. When this occurs, the male almost in 
a frenzy rushes on her, and they copulate with each of them embracing 
the other. 

Capture is carried out with the help of a large iron hook, one end 
of which looks like a ship's anchor (having a hook directed back- 
39 wards); while the other end has an iron ring to which a very long and 
strong rope is connected. A very strong man held this hook (haфoon 
V.H.) and sat with four— five persons aboard a boat. One of them 
handled the tiller while the other three or four rowed; and with them 
he carefully approached a whole herd of these animals. The hunter 

^^ According to the opinion of A.V. Vozzhinskaya (Institute of Oceanology of the 
Academy of Sciences), Steller wrote about the following species in order: Aganim sp.; 
Nereocystis sp.; Alaria ifistulosa); Zostera, or Laminaria longipes. 

^^Steller writes elsewhere that the intestine of these animals was always completely 
filled with large amounts of food: "The stomach is astonishingly large, its length is 6 
feet (180 cm) and width 5 feet (150 cm); it is filled with food and seaweeds to such 
a degree that four strong men with great difficulty can scarcely drag it from its place 
with a rope." 

The animal's feces "by its form, amount, odor, color as well as all other essential 
characteristics is recognized by the experienced stable-man as that of a horse". Steller, 
in his first days on the Island, considered these feces which were thrown up by the sea 
as horse feces, and considered this as a proof of his view, that America was not far from 
this place (no horses were present at that time in Kamchatka) (V.H.). 

-"". . . One of these animals was stranded on dry land because it slept and the sea 
at that time receded, so it was not able either to go back or to resist. It was killed with 
cudgels and axes. 



36 




Vyt.MM Ч,-' 



•fft "«. -»-л»* ■ 



ч 



^: : ^^^ ^^'0 ^^'^i- 



37 




■ .vJ- 






■JJf- 



36—37 Fig. 8. Bering Island. The coast on which Steller's sea cow was found. 

Photographs by S.V. Marakov. 



38 

stood on the anterior part of the boat holding the hook in hand; and 
when he was able to hit the animal, he did. Thirty persons who were 
standing on the shore holding the end of the cord tried to hold the 
animal and with great effort pulled it to the shore; the animal, at that 
time, resisted with fear. Those who were on the boat, held their place 
with the help of another rope, and continuously beat and stabbed with 
knives and other shaф instruments to such a degree that the weakened 
animal, completely exhausted and worn out by the wounds, was drawn 
up on the shore. 

Some [people] cut large pieces of meat from the animal while it was 
still living. During this, the animal only struck violently with its tail 
here and there, and so strongly resisted with its hands that large pieces 
of the superficial skin layer were usually detached from them. More- 
over, the animal was deeply breathing and moaning. Blood shot upwards 
as a fountain from the wounded back; so long as the head of the animal 
was hidden in water, there was no bleeding, and when it took a breath 
in air (for this reason it raised its head from water) bleeding began 



■■ >'^f-. ж 



^il^^_ ■ ■^»> 



- % 




Fig. 9. Peregrebnaya Bay near Cape Manati on Bering Island — the former habitat 
of Steller's sea cow. Photograph by S.V. Marakov. 



39 

again. This is because when the back was wounded, the lungs which 
He under it were also wounded and the more frequently the lungs were 
refilled with air, the more frequently they expelled blood .... 

The adult and very large animals are more easily hunted than 
calves, because the latter swim more rapidly and vigorously. If a har- 
poon was thrust in them [large ones] and was firmly attached to them, 
it remained in their bodies and was held there, but the calves sloughed 
off their skin and escaped, as was observed many times. 
40 If the harpooned and captured animal begins to struggle strongly, 

only the animals in the herd swimming nearby come close to it and try 
to help it. Thereupon some animals try to turn the boat upside down 
with their backs, while others lie on the rope^^ and try to cut it off or 
to strike the rope with their tails to pull the harpoon out of the back of 
the injured animal, and this they sometimes succeeded in achieving. 
Definite proof of their abilities and at the same time, of their conjugal 
love was shown in that one male whose female was captured 
by a harpoon and pulled out onto the sea shore, tried with all his strength 
to help her, but his attempts were unsuccessful, and in spite of all the 
blows which we dealt him, he followed the female to the shore, and 
several times suddenly threw himself like an arrow toward her although 
she was already dead. The next day, as we were getting ready early in 
the morning to cut the dead animal into pieces, we found the male still 
standing beside his female. Even on the third day, when I came to 
investigate the internal organs in detail, the male was still there. 

Concerning its voice, this animal is dumb and does not produce any 
sound. It only breathes loudly and also moans when wounded. But 
concerning sight and hearing, nothing can be said. These animals al- 
ways hold their heads under water and therefore they cannot hear or see 
well. This gives the impression that the animal itself has no need for 
these senses. 

. . . Several times it occurred that dead animals were cast up by 
storms around Cape Kronotskii, which is called the Nose, and Avacha 
Bay [on Kamchatka]. Because of the type of food which is consumed 
by these animals, the inhabitants of Kamchatka called these animals 
"Kapustnik" [cabbage-worm] in their language, which means eaters of 
weeds; I learned of this, when I returned (from Bering Island, V.H.) in 
1742. 



^^ According to the description of Bering's companion, Sofron Khitrov, rope of 4.5 
inch diameter, i.e. about 100-110 mm was employed (Pekarskii, 1869). 



40 

. . . The fat layer under the upper skin layer and the dermis 
covering the whole body is as thick as a palm's width, and in some 
places, three-quarters of its width. The fat is full of glands (!), soft, 
white in color, but in sunlight it turns yellow, like May oil. The odor 
and taste of the fat is pleasant and cannot be compared with that of any 
other marine animals, and it even exceeds that of tetrapod animals. 
Moreover, this fat can be kept for a long time even on very hot days, 
for it neither spoils nor smells. After mehing it is delicious and tasty, 
and is preferred to any other oil. Its taste is almost like sweet almond 
oil and it can be used for any purpose for which any oil is used. It bums 
brightly in lamps and produces no smoke or bad smell. . . The fat of 
the tail is harder, stiffer and therefore it is more delicious when cooked. 
The meat has stiffer and thicker fibers than beef; it is also redder in 
color than meat of terrestrial animals and it is suфrising that the meat 
of this animal can be kept in the very hot days for a long time without 
producing any smell, even if it is everywhere fiill of maggots[. . .у^ 

The meat of this animal, it is true, requires a longer time for 
cooking but when cooked, it has a superior taste and cannot be differ- 
entiated from beef. The fat of young sea cows is similar to that of fresh 
swine lard and it is hardly differentiated from the latter. Moreover, 
their meat cannot be differentiated at all from veal, and it also is 
cooked rapidly; during cooking, it also swells like young pork meat and 
occupies in the pot twice the space as originally. 

Fat in deposits, as in the head and tail, is viscid and therefore can 
scarcely be boiled. The most preferred muscles are those of the lower 
surface of the abdomen, back and flanks. Such tallow is not only fit for 
41 salting — ^which many persons doubted — but when salted it is like that 
of corned beef and has a very nice taste. Internal organs — ^heart, liver 
and kidneys — ^are very hard and we did not utilize them because we had 
enough meat, for a mature animal weighs nearly eight thousand pounds 
which equals eighty centners, br 200 Russian poods (3200 kg). The 
number of these animals was large on this single island, so the inhab- 
itants of Kamchatka might continuously feed only on them. 

A particular harmful insect*, like a louse, causes much trouble to 
sea cows. They stick in large masses on the wrinkled limbs, on mam- 
mary glands, teats, in hidden places, anus and in the shagreen-like 

^*The explanation given by Steller of this phenomenon is of пэ interest, but the fact 
itself of the possible long4listance transport of bodies by the sea, especially to Kamchatka, 
is significant (V.H.). 

*See footnote 19, p. 24.— Sci. Ed. 



41 

cavities of the upper skin layer. . . . These insects pierce both the upper 
layer of the skin and the dermis and protuberances are formed from the 
released fluid, and here and there visible warts. However, these insects 
are pursued by the sea-gulls which sit on the back of the animal and 
pick them up daintily. Thus they offer a pleasant service to the animals 
which suffer from these harmful insects . . .". 

Geographic Distribution and History of Extinction 

Steller discovered the kapustnik in 1741 on Bering Island. Later, it was 
established that this animal lived also on Mednyi [Copper] Island, but 
information about this is very scanty. In the historical period, sea cows 
were not found anywhere else. The sea cow is completely analogous to 
Steller's [Pallas's] cormorant {Fhalacrocorax perspicillatus Pall.), 
which was also limited, apparently, in its distribution to the Com- 
mander Islands and also was exterminated somewhat later — ^in the middle 
of the nineteenth century. 

It seemed that the sea cow must have been living in the Aleutian 
Islands, where the living conditions are actually the same as in the 
Commander Islands. The standing crop of sea weed is also great there, 
and the same species grow as in the Commander Islands; generally, 
they comprise one geographical and zoogeographical region. This par- 
ticularly applies to Attn Island, the most westerly of the Aleutians 
(Blizhniye group) and the nearest one to the Commander group (about 
350 km from Mednyi Island). 

Moreover, Steller did not refer to this species in places east of 
Bering Island in any of his works, including the diaries, that were 
published by Pallas (1793) after his death. There is also no record in 
the descriptions of any other voyages in the eighteenth century. As a 
result of a special collection of information and complete analysis of 
the question of the geographic distribution of this species, [von] Baer 
(1838)* noted the complete absence of this animal at any time in 
Russian America, including the Aleutian Islands, especially the Lis'ie 
and Andreyanov islands. 

Later, Stejneger (1883), in his evaluation of all information (tes- 
timony of Aleuts from Attu, data on Semitki and Agattu islands, as well 
as observations of Turner, Voznesenskii and others), denied the argu- 
ments, which are actually very vague, concerning the existence of 
Steller's sea cow on the western Aleutian Islands including Attu. 

♦Actually published in 1840— D.D. 



42 

He denies the existence of sea cows on Attn and [the report of] 
Nordenskiold (1882), based as it was on unwarranted dependence on 
testimony of inhabitants. A single rib found on Attn by Voznesenskii, 
if it actually belongs to this species, came from the body of a dead 
animal cast up by waves,^^ as Brandt (1846) correctly concluded. 

In American literature, including the latest (Murie, 1959; R. Rausch, 
1960), there is no evidence of the occurrence of Steller's sea cow 
42 around Alaska or on the Aleutian Islands, or any discovery of its 
remains (not taking into account the case given by Voznesenskii). 
There are neither archeological nor paleontological remains of this 
species in the American territories (information from Ch.A. Repenning, 
paleontologist, U.S. Geological Survey; only known are the remains of 
a closely related genus Halianassa, which is not younger than middle 
Pliocene, from the eastern part of the Pacific Ocean). In fact, if we 
leave aside possible historical causes, it is very difficult to imagine the 
emigration of a herbivorous animal which is so closely associated with 
the seashore and which has very little vagility, across 350 km of open 
ocean from Mednyi to Attn islands. 

Sea cows were never seen in Kamchatka and were generally absent 
on the Asiatic shores. Neither Steller (1751, 1753, 1781, 1793) nor 
Krasheninnikov (1755) made any reference to such an occurrence. All 
of Krasheninnikov's information (1755) was related to Commander 
[Bering] Island and, as mentioned above, was based on Steller's manu- 
scripts, which were at his disposal. There is direct information about 
the absence of this species in Kamchatka in the eighteenth century. The 
situation wherein some inhabitants of Kamchatka knew about this 
animal which they called kapustnik ("cabbage-worm") is explained by 
the washing up of bodies (see footnote on p. 40) and finding in their 
stomach after dissection of a large mass of "sea cabbage", as was cor- 
rectly noted by Stejneger (1883) (see footnote on p. 35; field cabbage 
was not known at that time in Kamchatka). The distance from the 
nearest point on Kamchatka (Cape Kamchatka) to Bering Island is 1 50 
km, a distance across which a carcass could be transported but cer- 
tainly insurmountable for a living animal. The absence of sea cows 
along the Asian coast was emphasized by [von] Baer (1838), who 
showed especially that this animal was absent not only in Kamchatka 
but also in the Kuril Islands. 

Later, Ditmar (1900) referred to the absence of sea cows on 
Kamchatka in his 1852 material. Finally, it is known that in the middle 

"This rib could have been brought by workers who were preparing meat in the 
Commander Islands (V.H.). 



43 



" , ■;^■•. iv,- 



,> .'■ , Jr . . ■ . 





Fig. 10. Tundra Bay on Bering Island — д^ place where sea cows occurred. 
Photograph by S.V. Marakov. 

of the eighteenth century, commercial expeditions to the Aleutian Is- 
lands and Alaska, for trade in beavers, [sea otters] and other fur-bearers, 
stopped off especially at Commander [Bering] Island to prepare sea 
43 cow meat and even wintered on the islands for this puфose. This was 
done because this animal was not found on Kamchatka (Pekarskii, 
1869; Ditmar, 1900 and others; see below). 

Some reports of the presence of sea cows on the Asiatic coast are 
found in the literature, but they are false. Thus, the reports of the 
observation (1762) of sea cows at Cape Shelagskii and between it and 
the Kolyma, in the ice of the East Siberian Sea (Grekov, 1958, from 
a document of Shalaurov), were based on some strange misunderstand- 
ing. Such a casual remark as Nordenskiold's (1882)* in which he says 
that "one Russian", around 1910, saw a dead sea cow supposedly 
washed up by the sea at Cape Chaplin (north of 64° N. Lat.) is not 
trustworthy. Finally, in the light of what has been said above, and as 
all the data on the biology of the sea cow make absolutely clear, 
"observations" of this species at Cape Navarin in the Bering Sea (about 



•= Actually attributable to Sverdrup (1930) — D.D. 



44 

62° N. Lat.) in 1962 (Berzin, Tikhomirov, Troinin, 1963) have no 
value at all (Heptner, 1965, see below). 

The history of the extinction of the sea cows is considered the most 
tragic history of the destruction of a most remarkable animal in a very 
short period, and at a time when the technology of animal destruction 
was at a low level. Steller's sea cows were in the true sense of the word 
eaten up by human beings. 

As is clear from Steller's description and from other information 
(see below), the meat and fat of sea cows were characterized by their 
excellent quality. This fact was sufficiently widely advertised by Steller, 
who returned to Kamchatka in autumn 1742. Moreover, he suggested — 
this probably was a mistake — that the number of sea cows around the 
island was very large. In fact, these animals were, apparently, numer- 
ous only where the crew of the "Saint Peter" was shipwrecked and 
spent the winter, and in some other especially favorable parts of the 
coast. The total number of these animals could hardly have exceeded 
some hundreds. 

What is then the cause for such intensive destruction of this animal, 
which lived around the shores of two similar, small islands lying far out 
in the ocean, and which had been just discovered and were uninhab- 
ited? On the one hand, Steller wrote diligently about the quality of the 
meat and fat of the sea cow, and especially about the fact that Kamchatka 
at that time could not supply with foodstuffs the numerous commercial 
ships which were sailing from Petropavlovsk and Bol'sheretsk in 
Kamchatka to the east and northeast. The Commander Islands therefore 
became a base for obtaining provisions for the crews of these ships. 
The system of hunting was constructed suфrisingly rapidly and its 
organization was very flexible, and the "profusion" of the island was 
rapidly appropriated. In the next year after Steller's return, i.e. in 1743, 
the ships which were sailing to trade in the Aleutian Islands and in 
Russian America went at first to Bering ("Komandor") Island and 
worked there preparing sea cow meat, overwintering because of this. 
Later this was continued by all the numerous commercial ships directed 
from Kamchatka to the east for hunting arctic foxes, fur seals and, 
above all, Kamchatka beavers [sea otters]. 

The Marine Department in Petersburg in 1765, apparently in ques- 
tioning the merchant Vasilii Shilov about the distant lands situated 
beyond the Commander Islands, and about the trade and geographical 
discoveries conducted there by the commercial companies, asked him 
particularly about the "means" that they had in those far and long 



45 

voyages ". . . how you were saved from hunger, because the food 
prepared in Kamchatka was for two or three months, and flour was 
especially insufficient?" The answer declared that the "trading compa- 
nies at present are far from Kamchatka beyond Commander and Mednyi 
and near to the North, and of others, several islands became better 
known far to the East", however, Kamchatka "cannot provide such 
distant routes with provisions, especially for prolonged residence there 
44 for hunting, and thus they used, except for flour, the abundance of 
Commander and Mednyi islands". From Kamchatka, the ships took 
besides flour, only such provisions and fresh water as helped them, 
even in unfavorable weather, to reach the Commander Islands ". . . and 
there they spend the winter; and during the whole wintertime they are 
engaged in hunting, and on the distant routes they prepare provisions 
which mainly consisted of sea cows, whose meat is not only wholesome 
and in great quantities, but can also be used instead of bread when 
dried. In this animal there is much fat to be stocked in small barrels and 
used in meals instead of oil, and one can drink it harmlessly like the 
best [olive] oil. It can also be used for making fire instead of candles. 
The skin of these sea cows is used instead of boards in the planking 
of boats . . ." (Pekarskii, 1869). 

Hunting itself was, on the whole, performed in the same way as 
that of the people on Bering Island, and it was based on the same 
biological characters of the animals, i.e., on their habit of coming to the 
shallowest water at ebb tide. However, some changes appeared in the 
technique. Peter Yakovlev describes it in the Nizovtsovaya harbor, 
where ". . . in the sea, near the shore, these sea cows were seen singly 
or in groups. In hunting they go in a boat, eight in a group, of which 
one hunter stays near the stem in front of the helmsman holding a long 
pole armed with a wide, long sword-like band (called "pokolyuga"), 
and the rest of the hunters sit as oarsmen. When they find themselves 
in front of a sea cow's head, they turn the boat with the stem towards 
the sea cow and the hunter with the "pokolyuga" inflicts a severe 
wound in that beast's heart. After that the rowers must row quickly 
away from the sea cow in order to get out of that wounded sea cow's 
reach, for it can min their boat by its tail or fluke . . . with such a 
wound in its heart, the cow does not swim in the sea for a long time. 
It soon gets tired, stops and turns with its belly up. The hunters pull that 
sea cow by a hook [haфoon, V.H.] towards their boat, tie it to the stem 
of the boat and row to the shore. The meat of one sea cow was enough 



46 

for 33 men for a month. The net weight of one sea cow's meat and fat, 
without bones, reached 200 poods^^ (Pekarskii, 1867). 

Moreover, as Vasilii Shilov describes, workers used the skins of 
sea cows in covering the frameworks of boats instead of covering them 
with wooden boards. From Kamchatka, they took "the ready-made 
wooden boat keels and the stem and bow installations for one or two 
boats" and covered them with sea cows' skins in the Commander 
Islands, "because" he notes, "such large skins are not present in 
Kamchatka" (this also proves the absence of the above-mentioned species 
on the continent). Sewn skins (that is, from animals of smaller size) are 
not convenient because they leak. 

Shilov further stated that "though the ship-chandlers take with 
them all the prefabricated parts, until they are actually needed (almost, 
V.H.) they do not cover them with the skins but prepare excess skins 
(my emphasis, V.H.). Nevertheless, they store up skins in a great 
quantity and when necessary they gathered the boats in a short time, 
and covered them with skins after soaking them in water . . . ." These 
skins were, apparently, of a great value because the boats were safer 
in the breakers, and what is more important, they were comfortable in 
handling and very light in comparison to wood-planked ones. "A boat 
covered with leather carried the same number of people (as a wood- 
45 planked one — ^20 men, V.H.) and it could be pulled out onto the shore 
or from the breakers, like a bladder, without risk, and could be carried 
a good distance by only four men. ..." 

As shown, Steller's sea cow was used by pioneers in developing 
Kamchatka and Russian America rather fully, but too excessively. As 
a result, it was quickly exterminated and in 1768, that is only 27 years 
after its discovery, the last one was killed on Bering Island. The ani- 
mals around Mednyi Island, which were, undoubtedly, fewer, were 
exterminated earlier; at any rate, in 1754 there was not a single one of 
them.'^^ It is possible that on Mednyi Island one or two outlived the 
indicated date for a year or two; however, in the seventieth decade of 
the eighteenth century nobody saw them and Bragin, who was spending 
the winter on Bering Island in 1773—1774, makes no mention of sea 
cows. There are no later records of them (Berg, 1946). 



^^Steller (see above) apparently considered that 200 poods is the total weight of the 
beast (V.H.). 

^'Peter Yakovlev, sent by the government according to the suggestion of M.V. 
Lomonosov, to Mednyi Island to search for useful minerals (copper), whose whole 



47 

The rapid extermination of sea cows was caused by thoughtless, 
rapacious forms of exploitation which led to the unjustified annihilation 
of a great number of animals. As seen from the descriptions of the 
methods of hunting, success was only ensured in cases where many 
people at once (not less than 20) were hunting one animal. Attempts to 
get an animal by a small group of hunters were often useless. The 
above-mentioned P. Yakovlev writes of this as follows: ". . . those 
hunters live in yurts or winter houses in different places along the 
northern shore of the Commander Islands in groups of 2 and 3 hunters 
and they hunt for arctic foxes and other marine animals. They do not 
have any other food except the meat of [sea] cows; the herds of cows 
live in the sea near the shore and are frequently wounded by people, 
but those wounded cows swim out to the sea and there become further 
weakened by their wounds. They are later washed ashore after a long 
period of time when their meat becomes bad and inedible. 

By this sort of hunting sea cows are eradicated, and although many 
are slaughtered (my emphasis, V.H.), not a single fresh sea cow comes 
to their hand, and the hunters therefore suffered from starvation, and 
the sea cows were exterminated." Yakovlev foresaw the quick exter- 
mination of the sea cows on Bering Island and tried to take measures 
for the protection and wise use of the stocks of this most important 
animal at that time. "For this puфose, on his (Yakovlev' s) return from 
one sea voyage (i.e. sailing to the Commander Islands, V.H.) it was 
reported on the 27th of November, 1765* to the Nizhne-Kamchatskii 
Ostrog [fort] that hunting of the sea cows should be prohibited by order 
(Dokladnaya Zapiska, V.H.) of the Bolsheretsk office in order to save 
one Commander Island [Bering] from devastation because Mednyi 
Island had already been devastated . . ." (Pekarskii, 1867). However, 
as is known, nothing came of Yakoylev's attempt. In this way, the sea 



expedition consisted of 33 persons, could not spend the winter on this island in 1754,** 
i.e., 13 years after the discovery of sea cows on Bering Island, because sea cows were 
absent .... "Mednyi Island was found empty — he wrote — and ... for the expeditionary 
force to spend the winter was impossible — due to the absence of sea cows the inhab- 
itants of this island suffered from hunger." Further, he wrote that "before the sea cows 
were found in large herds but now these cows were destroyed, so none of them could 
be seen,- and now in the Mednyi Island there is not a single sea cow . . ." (Pekarskii, 
1 867). Yakovlev and his "command" spent the winter on Bering ("Commander") Island 
(see below). 

*This date should be 1755, since 13 years later, in 1768, the last one was killed 
on Bering Island (see previous page) — ^D.D. 

**Misprinted 1745 in Russian original — ^D.D. 



48 

46 COW at Mednyi Island was destroyed ten or twelve years after 
its discovery. After approximately the same period, they stood under 
threat of extermination at Bering Island, and 13 years following 
Yakovlev's attempt to protect this animal, was also completely 
destroyed there.* 

All information about the existence of the sea cow, which occa- 
sionally appeared in the literature at the end of the eighteenth and the 
beginning of the nineteenth century, as was shown by K. [von] Baer 
(1838) and J.F. Brandt (1846), cannot be credited. Separate informa- 
tion appeared later concerning some of these animals (for some, see 
Grekov, 1958). The most famous of such reports, which seems most 
plausible, is that of Nordenskiold (1882), in which it was mentioned 
that a sea cow was observed on Bering Island in 1854. As was ex- 
plained by Stejneger (1883, 1885), who interviewed the same people 
to whom Nordenskiold referred, this information was found to be false. 
Stejneger (1883) suggested that these animals were whales of smaller 
size or, probably, female narwhals, which very rarely passed so far to 
the south and therefore were considered unusual for the Aleutians. 
Some other unreliable and unconfirmed information is not likely at all 
and is not worth discussing. Some of it is simply fantastic (observations 
in . . . California!). 

Even the slightest probability is lacking in the latest information 
given by Berzin, Tikhomirov and Troinin (1963), on the observation of 
a sea cow in July 1962 at Cape Navarin (!), i.e. somewhat north of 62° 
N. Lat. This information contradicts all the known facts about this 
species, which are partially given above. It was based on complete 
ignorance of the question and its literature (even the new literature), on 
the biology of the animal, characteristics of its habitat, and its behavior 
as well as the history of discovery and exploitation of northeastern 
Siberia and Russian America in the eighteenth and nineteenth centuries. 
This was actually the result of naive pursuit of sensation. 

Still, in our time, remains of sea cows are sometimes found. These 
remains have become rare, although a skull was found in the decade of 
the 30's and two lower jaws in a well-preserved condition were col- 
lected in 1960 (N.N. Kartashev, Zoological Museum of Moscow 
University). 

There are, in general, numerous bones and skeletons of sea cows 
preserved in museums of our country and in foreign museums which 

*See foot note on previous page — Sci. Ed. 



49 

were collected from the Commander Islands. It is interesting that at the 
beginning of the last century, a masticating plate was the only material 
at the disposal of [von] Baer, and Brandt ( 1 846) had only one imperfect 
skull. At the beginning of the 80's, Stejneger (1883) collected more 
than 1 1 skulls and believed that more skulls could be collected. B. 
Dybovskii brought some specimens from the Commanders in those 
years. Now, such collections are impossible* (V.H.). 



"Specimens have been collected as recently as 1992 — D.D. 



01ШЕК OF CARNIVORES 
Ordo CARNIVORA Bowdich, 1821 



Superorder of Carnivores 
Superordo FERAE Linnaeus, 1758 

Order of Carnivores 
Ordo CARNIVORA Bowdich, 1821 



49 The order includes species of relatively generalized type and clearly, 
sometimes very sharply, specialized ones. A few are large, the majority 
are of middle and small size, while some are of very small size. They 
are typically terrestrial animals, in few cases, semiaquatic, and as an 
exception aquatic animals of various construction and appearance, from 
heavy and clumsy animals to light, slender and very mobile. The tail 
is usually long (about half the body length or more), rarely short, of 
approximately the length of the hind foot. 

The extremities are plantigrade, semidigitigrade or digitigrade, digits 
are usually 5, but in some one is absent. Digits are provided with 
claws, in some species retractile; the terminal phalanges are not flat- 
tened and not widened, but always laterally compressed. One digit is 
never opposed to the others. In the wrist, the scaphoid (scaphoideum), 
semilunar (lunatum) and central (centrale) bones are fused. The articu- 
lating surface of the astragalus is pressed inward. The ulna and fibula 
are normally developed and separate (not fused).* The foramen 
entepicondyloideum on the humerus is either developed or absent; the 
third trochanter of the femur is absent. The clavicle is either absent or 
rudimentary. 

There are no more than 22 lumbothoracic vertebrae, usually 13 
thoracic, and 7 lumbar. The vertebral column is, in the majority of 
cases, very flexible. The number of caudal vertebrae varies greatly. 

The form of the skull is extremely varied — ^in some animals it is 
elongated with highly developed facial part and a relatively short cra- 
nial one, while in others the skull is very short, with rounded outline 
and with relatively large and voluminous cranium. All the transitional 
forms are also present. The hard palate is continuous, the eye socket is 
usually open, and only exceptionally superficially separated, but in the 

*Sic; refers to radius-ulna and tibia-fibula being distinct bones — Sci Ed. 



54 

majority, it communicates widely with the temporal fossa. The eye 
sockets are directed forward and in several, strongly approximate 
(binocular vision). The tympanic bulla (with one exception — Nandinid) 
is ossified. The wing-shaped (ectopterygoid) fossa (f. pterygoidea, s. 
ectopterygoidea) is not manifested in the majority of cases due to the 
absence of the corresponding independent process of alisphenoid. 
50 In the greater number of species, the crests and zygomatic arches 
are strong and widely separated, a fact associated with the great devel- 
opment of the jaw musculature. Ethmoturbinal (ecto- and endoturbinals) 
olfactory folds are 5-7; the maxilloturbinal has double flexures or branch- 
ing. The articular condyle of the lower jaw is semi-cylindrical and the 
glenoid fossa on the skull is elongated, occupying a transverse position. 
It is usually more or less deep, sometimes so much so that its anterior 
and posterior edges tightly envelop the condyle of the lower jaw. The 
angular process of the lower jaw is small and with no inward flexure. 
The dental system is heterodont and diphyodont. The complete 

3 14 4 
formula is I— C-P — M— = 48, but usually the teeth are much fewer. 
3 14 4 ^ 

The reduction occurs at the expense of the cheek teeth and mainly at 

the expense of the posterior molars and (to a lesser degree) of the 

anterior premolars. The number of incisors is only rarely reduced to 

3 2 

— (Kamchatka "beaver" or sea otter,* Enhydra) or to — (sloth bear, 

3 12 1 
Melursus). The minimum dental formula isl— C-P — M- = 28 (manul, 

3 12 1 

3 111 
Felis manul) and even I— C-P — M- = 26 (fossil sabre-tooth tiger, 

3 12 1 

Smilodon, of Machairodontinae). 

In one case (bat-eared fox,** Otocyon) the dental formula may be 

3 1 4 3-4 
I-C-P-M^^-— = 46-48-50 ' 

3 1 4 4-5 

*ln Russian, Kalan — Sci. Ed. 

**In Russian, long-eared fox — Sci. Ed. 

'The greatest number among recent placentals. Probably, this increase is a 
secondary character and is connected with the transition to a considerable extent to 
entomophagy (mainly grasshoppers). 



55 

The teeth always are rooted, brachyodont, cusped-cutting (tuberculo- 
sectorial), quadri- or multituberculate with shaф, or rarely blunt, apices. 
The teeth are strongly differentiated. Incisors are small and the canine 
is usually strongly developed. The fourth upper premolar and the first 
lower molar are usually large, with shaф cutting apices ("camassial 
teeth"), rarely with a more or less flat, bluntly tuberculated, mastica- 
tory surface. A true diastema is absent. 

The stomach is simple; the caecum is normally or weakly devel- 
oped or absent. The brain is macrosmotic, the hemispheres are large 
with three suprasylvan fissures. The pelage is always well developed, 
composed of different hair categories (underfur, guard hair, directional 
hairs), sometimes very dense, relatively long and fluffy, and sometimes 
sparse and rough. The tail is usually fluffy. Vibrissae are well devel- 
oped. The color varies greatly, being more or less monotone, or light 
with bands, spots, shabrack*, etc.; some species are polymorphic. The 
skin glands, with some exceptions, are normally developed; the major- 
ity of glandular complexes and large, complicated, often odiferous glands 
(musky stink glands of skunks, etc.) develop in the anal region in 
connection with the male genitalia. The milk glands — ^inguinal and ab- 
dominal — range from 6-7 to 2 and one pairs. The scrotum is well 
developed or else the testes are located subintegumentarily, but always 
outside the abdominal cavity ("extra-abdominal"), and behind the copu- 
latory organ. In the latter (except in hyena) a bone (os penis, baculum**) 
is present. The uterus is doubled or with 2 horns. The placenta is 
deciduous and zonaF. 

Sexual dimoфhism is absent or weakly manifested except in the 
general size of the species with rare exceptions (lion). Age dimorphism 
in some species is quite significant (wolf, lion and others), while in the 
51 rest it is not manifested. Seasonal dimoфhism, mainly in the density 
and length of fur in the forms of temperate and northern latitudes, is 
quite great and sometimes connected with changes in color, in which 
case it is shaф (arctic fox, ermine), while in the others it is weak or 
absent. In the forms of temperate and cold latitudes that are active in 
winter, there are two molts which usually occur rapidly. In species of 
those districts which spend the winter sleeping, the single molt extends 

*Lit., goatskin; refers to a goatskin saddlecloth, hence by extension, to a "saddle" 
pattern on the back — Sci. Ed. 

** Spelled bacculum in Russian original — Sci. Ed. 

^Occasional reference to the placenta as of the discoid type (for example, Novikov, 
1956) is a mistake. This was, apparently, based on a slip of pen in the diagnosis of the 
order by Weber (1928). 



56 

over almost the whole summer (the summer fur is considered a tran- 
sient coat). In warm countries, there is usually one molt. 

Carnivores differ greatly in size — the largest has a head-body length 
(excluding tail) to 300 cm and a weight to 725 kg and even 1 metric 
ton (polar bear). The smallest form — the least weasel {Mustela nivalis 
pygmaea) has a body length of 1 15— 140 mm and a weight of about 
100 g; i.e. the smallest weight is less than the greatest one by about 
10,000 times (in case of artiodactyls this proportion is about 1 : 1,150). 

In general constitution, proportions and external appearance, bio- 
logical types, adaptation forms and other characteristics, the carnivores 
are one of the most variable if not the most variable, of the orders of 
mammals, including forms that are comparatively generalized types 
(some bears) as well as extremely specialized ones (Kamchatka bea- 
ver, or sea otter). 

Among the carnivores there are heavy animals of low mobility and 
slow movement (bears) as well as light and slender, rapid and swift 
runners (wolf, cheetah), excellent swimming forms closely connected 
with water (otters) and excellent climbers, and even partially special- 
ized arboreal animals (some martens, viverrids and cats); some are 
excellent diggers (badger), etc. Carnivores inhabit all latitudes, land- 
scapes and the vertical montane belts. They live individually or in 
pairs, and some are found in small packs (wolves, hunting dogs) at least 
in part of the year. The majority are sedentary and are fairly strictly 
restricted to their hunting areas; others wander widely, while several 
accomplish regular seasonal migrations. 

Some species are associated the entire year with holes (badger); 
others, only in some parts of the year (time of reproduction) and a third 
group has only open lairs even in the lactation period. Some are asso- 
ciated continuously, or only at the time of reproduction, with tree 
hollows, rock clefts, caverns and other similar shelters. Several species 
of cold and temperate latitudes hibernate in winter. A series of species 
are monogamous and live in pairs for more than one reproductive 
season, while others form pairs only for one cycle and in others, the 
male does not take any part in provisioning the young at all. True 
polygamy (in the form of a "harem") does not occur. 

The young range from 1-2 to 13 and even 20-22.^ However, the 
number of yo\mg in the litter varies strongly within one species. The 

^Maximal known number is in the wolf (13) and in arctic fox (22). Normally, these 
numbers are much less. 



57 

offspring are bom helpless, blind with closed ear openings and undi- 
vided fingers, etc. and they develop and grow comparatively slowly. 

Carnivores are extremely variable in their food habits. The major- 
ity are sarcophagous, at times very narrowly specialized, feeding on 
the meat of mammals and birds, and rarely ichthyophagous and polypha- 
gous, in the diets of which a significant role is played by lower 
vertebrates, terrestrial and aquatic (mainly fresh water) invertebrates 
and plants. The number of species that are chiefly plant-eaters 
(phytophages) is few; specialized entomophages comprise exceptions, 
(aardwolf* — Proteles). 

Among the sense organs, smell and hearing are well developed, 
sight is usually weak, and not color. The external ear pinnae are very 
different in size, from equal to head length (fennec) to almost undevel- 
oped (otter). 

Carnivores are very widely distributed, occupying nearly the whole 
53 world. Aboriginal species are absent only in Australia and in some 
islands and island groups. In the north, they are met with in the north- 
em most islands (New Siberian, Sevemaya Zemlya, Franz Joseph Land, 
Arctic archipelago of [North] America, Greenland) and even in the ice 
of the Arctic Ocean, considerably to the north of these islands, all the 
way to the pole (polar bear, arctic fox). The species of other orders, 
even the pinnipeds, do not penetrate so far north. Camivores are absent 
on the island of the West Indies and the other islands of the Atlantic, 
except for Newfoundland, Iceland and Jan Mayen. They are also ab- 
sent, with the exception of the Falklands, on the islands of the southern 
oceans, small islands of the Indian Ocean, in Tasmania, New Zealand 
and New Guinea, Vetar [Wetar] and Ombai, in the Lesser Sundas (on 
others, including Timor in the east, they are present), Tenimber 
[Tanimbar], Kei and the majority of the Moluccas (except for 
Halmahera, Bum and Ceram), in the Talaut [Talaud] archipelago and 
on all the islands which lie to the east of New Guinea and on the 
Hawaiian and Galapagos Islands. They are found in Ara, Sulawesi 
(Celebes), Philippines, Ryukyu, Kuril, Sakhalin, Commander, Aleutian 
and Pribilof islands. 

The geographic range of the order has not essentially been changed 
since historical time and it was not reduced — ^with some exceptions 
[extiфation of the Falkland "wolf, Canis {Dusicyon) australis 
= antarcticus]. The geographic range was artificially widened for some 

*In Russian original, maned "wolf — Sci. Ed. 



59 

species; the Indian mongoose was established in some islands of the 
West Indies, Fiji islands and several others; the European red fox was 
introduced into Australia, etc. The dingo ("Canis familiaris dingo"), 
occupying the entire continent of Australia, is not an aboriginal species. 
This is a feral dog, which was introduced into Australia a very long 
time ago by the ancestors of the contemporary Australians as a domes- 
tic animal."* 

On account of the diversity of biological types and forms of spe- 
cialization which allow carnivores to occupy almost all habitable media, 
the breadth of their geographic distribution, the diversity and abun- 
dance of species, the relatively high numbers of many of them, and 
other characters, carnivores without doubt may be considered a flour- 
ishing and quite recent group. Although anthropological factors have 
negatively influenced a series of species, decreasing their geographic 
ranges and numbers, this influence is, however, generally less in rela- 
tion to this order than if compared to several others. The "resistance" 
of carnivores to this very strong destructive factor is an indication of 
their vitality. 

Concerning systematic relationships, carnivores are closely related 
to pinnipeds (Pinnipedia). The latter are considered to be derived di- 
rectly from carnivores comparatively recently (Oligocene). All their 
characteristics have been formed as shaф and direct progressive adap- 
tations to an aquatic (marine) way of life. Some more primitive recent 
pinnipeds (Otariidae — eared seals) are, in their least specialized charac- 
ters (skull), very similar to the generalized forms of Camivora (Steller's 
sea lion,* Eumetopias). Many authors unite carnivores and pinnipeds 
in a single order Camivora, dividing both groups only into the rank of 
suborders: Fissipedia and Pinnipedia. Such a view was mainly based on 
paleontological data. The alignment of the recent groups (by the "hori- 
zontal" system), however, is such that for the purpose of keeping with 
the sequence in the mammalian system, it is more natural to consider 
Camivora and Pinnipedia as separate orders.^ 



"•The origin of the dingo from the primitive dogs of the tropical zone (Shensi type 
"Canis familiaris schensC) is indicated by the fact that the species of the genus Canis 
are absent in the Malayan archipelago, and they (wolves and jackals) begin to occur only 
in Indo-China and India. 

*In Russian, sivuch — Sci. Ed. 

^Accordingly, in this edition, the suborder Fissipedia is opposed, not to the subor- 
der Pinnipedia, but to the suborder Creodonta. In Simpson's system (1945), Creodonta, 
Fissipedia and Pinnipedia are considered as suborders of the order Camivora. 



60 

54 The features that approximate carnivores with insectivores are 
relatively great, and those which link them with artiodactyls and peris- 
sodactyls are strongly divergent, and essentially in a purely historical 
relationship (see vol. I). Their relationship with whales, which is fre- 
quently emphasized and sometimes regarded as very close, is apparently 
not more close than the relationship to ungulates. 

The ancient and most primitive group of Camivora — Creodonta — has 
a great phylogenetic significance. They stand at the root of the exten- 
sive cohort Ferungulata, comprising the orders of pinnipeds (Pinnipedia), 
tubulidentates (Tubulidentata), probiscideans (Proboscidea), hyraxes 
(Hyracoidea), sirenians (Sirenia), even- and odd-toed ungulates 
(Artiodactyla, Perissodactyla) (see p. xx). 

Camivora is one of the most ancient orders of recent mammals, 
stemming indirectly from the insectivores, the most primitive and an- 
cient placentals. The primitive carnivores, the lower Creodonta 
(Arctocyonidae), appear in lower Paleocene (North America) and are 
represented by forms which are intimately connected — in certain re- 
spects "transitional" — with primitive Cretaceous insectivores 
(Insectivora). Creodonts lived through their flowering in the Eocene 
and were still relatively well, but notably less richly developed, in the 
Oligocene. Their last representatives (Hyaenodontinae) survived only 
to the very beginning of the Miocene. True Creodonta were represented 
by several families (Arctocyonidae, Mesonychidae, Oxyaenidae, 
Hyaenodontidae) which did not leave any trace. On the same level with 
these families of the typical creodonts, the family Miacidae existed 
from the middle Paleocene (became extinct in the Eocene), which by 
general agreement, was the beginning of modem camivores (Fissipedia). 
The unique features of this family — ^which are, to a certain extent, of 
"intermediate" character between Creodonta and Fissipedia — gave some 
authors the basis for considering them among the Creodonta and gave 
others the basis to relate them to the existing Camivora and to ally 
them to viverrids (Viverridae), or sometimes to cats (Felidae) or to 
wolves (Canidae). If the Miacidae are considered among the existing 
camivores (Fissipedia), then the origin of this latter group (from the 
primitive Creodonta) must be dated to the middle Paleocene. The re- 
maining (recent) families of tme camivores (Fissipedia) appear not 
earlier than the upper Eocene and the lower Oligocene. 

Among the sum of characters, the contrast between the extinct and 
recent groups of camivores is relatively small, although in the extinct, 
even a separate suborder is recognized (see below). It is less than, for 



61 

example, in the "ungulate" group. The diversity of the group in the past 
was similarly not greater than in the more recent (and present) time. 
The same applies to the general number of species in different preced- 
ing epochs. Therefore, in its historical aspects there is basis for 
considering the order of carnivores as an existing flourishing group. 

Classification of the order of carnivores, in spite of the abundance 
and diversity of species, is, ii: its main features, simple and does not 
result in any serious disagreements. Two suborders are admitted — the 
extinct Creodonta with four families (see above) and the recent 
Fissipedia. The latter is divided into three superfamilies: the extinct 
Miacoidae with one family, Miacidae; the recent Canoidea (Arctoidea 
auct.) with four families — Canidae (wolves), Ursidae (bears), 
Procyonidae (raccoons), Mustelidae (martens) and the recent Feloidea 
(Aeluroidea, Heфestoidae auct.) with three families — ^Viverridae (viver- 
rids), Hyaenidae (hyaenas), Felidae (cats).^ 
55 Canoidea are characterized by the following characters. 
Tympanicum formed by the whole outer wall of auditory bulla. Parti- 
tions of internal cavity of auditory bulla derived from cartilagenous 
precursors, or precursors of the tympanicum. Canalis caroticus long. 
Maxilloturbinalia large and branching, and partition off the nasoturbinalia 
and ethmoturbinalia from the external nasal openings, Cowper's glands 
absent. Os penis large. 

Feloidea are characterized by the following characters. Tympanicum 
is, to some degree or other, semicircular and forms only the anterior 
outer (tympanic) part of the auditory bulla, while the posterior inner 
part is formed by the entotympanicum (os bullae). The tympanal- 
entotympanal partition extends deep inside the bulla at the place of 
divergence of both bones. Canalis caroticus short or absent. 
Maxilloturbinalia small, doubly flexed, and do not partition off the 
nasoturbinalia and ethmoturbinalia, which, in this way, approach the 
external opening of the nasal passage. Cowper's glands present. Os 
penis small or absent. 

Canoidea and Feloidea are sometimes considered to be suborders. 

The systematics of carnivores is insufficiently worked out as re- 
gards the genera and species. In particular, there is a strong tendency 

^Attempts were often made to divide some families into several. Among Procyonidae 
in particular, the [lesser] panda {Ailunis — ^Ailuridae) and giant panda* {Ailuropoda — 
Ailuropodidae) were divided into separate monotypic families. These, and also other 
arrangements, cannot be considered well-founded and are not widely accepted. 

*In Russian, bamboo bear — Sci. Ed. 



62 

here towards narrow 1п1ефге1а11оп of the genus (especially cats, bears 
and some others) and species (particularly in America).^ According to 
a more or less moderate opinion (Simpson, 1945), the order comprises 
338 genera belonging to 12 families, of which 5 are extinct. Extinct 
genera number 241 ; recent — 91} 

Only a very approximate count can be made concerning the num- 
ber of the recent species of carnivores at the present moment. Their 
general number is approximately 240 (probably less), constituting only 
8% of all existing mammals. On the average, the genus comprises 2.5 
species. The greatest number of species is in the families Viverridae 
(about 75) and Mustelidae (about 70); in Canidae there are 29, Felidae — 
about 36, in Procyonidae — ^about 10, Ursidae — 1, and Hyaenidae — 4. 
The greatest number of species live in Africa (about 100) and Eurasia 
(about 90); in South America — ^about 50 species, in North America — 
about 35.' 

In practical relationships, carnivorous animals are one of the most 
important groups of mammals. Among them, there are a great number 
of fur-bearing species, some of the most valuable (sea otter, sable), 
pests of livestock and hunting enteфrises (wolf, raccoon dog, yellow- 
throated marten), destroyers and regulators of a number of harmfiil 
animals (mainly rodent — ^foxes, some martens, viverrids and others), 
valuable sport-hunting animals (large species of cats, bears), vectors of 
some domestic animal and human diseases (rabies), animals giving raw 
material for perfumes (musk), etc. 

Some species have been acclimatized in alien countries as fur-bear- 
ing animals (raccoon dogs in European part of the USSR, American 
raccoon, American mink in the USSR), as destroyers of harmful ani- 
mals (snakes in the West Indies — mongooses). However, these measures 
are not widely distributed in the sense of territory and number of species. 
A series of species, as a result of the excessive utilization in connection 
with the value of their fur or pursued for the sake of destruction for one 
56 or another reason, sometimes very strongly, were reduced in number 

'Lately, a wider approach to the species and genus concepts has been applied in 
the systematics of carnivores, and it is well-known to be found in a revisionary stage 
(see below in characters of families). 

"The relatively high "index of contemporaneity" of carnivores indicates that con- 
temporary genera constitute a sufficiently large part of the total number of genera — more 
than 28%. 

'According to Anthony, 1928; Allen, 1939; Cabrera, 1957-1961; Chasen, 1940; 
Carter, Hill and Tate, 1945; Anderson, 1946; Ellerman and Morrison-Scott, 1951; 
Roberts, 1951; Haltenorth and Trense, 1956; Haltenorth, 1958 and other sources. 



63 

and distiibution (sea otter, lion, tiger, bears, wolf, leopard, ршпа and 
others). Some forms are exterminated completely or almost completely 
(Falkland wolf, Cis-Caucasian leopard, Persian and Atlas lion and 
others), A series of species find themselves under a complete or partial 
[protective] ban, and some others are kept in preserves. The hunting of 
the majority of fur-bearing and sport-hunting animals is regulated. 
Conservation of carnivores usually gives good results. 

Carnivores gave rise to several domestic animals. The most impor- 
tant of these is the dog ("Canis familiaris") which is the most ancient 
of all the domestic animals, derived from the wolf (Ca«/5 lupus)}^ The 
domestic cat ("Felis catus") is a minor, mainly decorative, domestic 
animal, obtained as a result of the domestication of Felis sylvestris 
libyca. The white polecat {Mustela putorius furo) — the albinistic form 
of the polecat — ^is only a semidomestic form from western Europe. In 
the last decades, a series of species (red fox, Vulpes vulpes — melanis- 
tic form; Arctic fox, Alopex lagopus—nonnal and "blue" forms; 
American mink, Mustela vison; raccoon, Procyon lotor; sable, Mustela 
zibellina, and several others) became the objects of different forms of 
fiir farming, mainly hutch farming, and many different breeds of foxes 
or minks have been derived. 

In the fauna of USSR there are 41 autochthonous and 2 exotic 
acclimatized species, i.e. about 17% of the species of the order, and 
about 14% of the species of the USSR fauna. Both superfamilies, 5 out 
of the 7 families (71%) and if the acclimatized American raccoon is 
considered, 6 families (85%), and 22 genera out of the 97 existing ones 
(22.5%) are present in the USSR." 

The geographic range of carnivores in the USSR covers the whole 
territory of the country and the Soviet sector of Arctic Ocean (they are 
now exterminated in the southwestern part). 

In the USSR, representatives of the order are found in all natural 
conditions, including extremes of high mountains, deserts and the sea- 
shore. 

In the Soviet Union, carnivores constitute a very important group 
practically. They include the most valuable fur-bearing animals, some 
of them represent objects of hunting sports, while certain species play 
a more or less considerable role as pests of livestock and hunting 

'"The presumption of the participation of jackal, i.e. the diphyietic origin of the dog, 
is to the highest degree improbable and is now entirely rejected. 

"The number of genera, also mentioned above, is that given by Simpson (1945). 
Later in the text, several genera will be discussed in a broader manner. 



64 

еШефпзез, carriers of some diseases, and destroyers and regulators of 
the numbers of harmful rodents. The hunting of almost all species is 
regulated (definite times of hunting, licensed trade); some are found 
under complete or partial protection, while certain species are com- 
pletely exterminated from the country or from some parts of it (V.H.). 

Key for Identification of Families of the Order of 
Carnivores 

(Identification by External Features) 

1 ( 6). Five digits on hind foot. 

2 ( 3). Tail short (shorter than the hind foot) and hidden in body fur 

or hardly protrudes from it. Size very large, body length of 
aduh 1.4-2 m or more Bear family, Ursidae (p. 586). 

3 ( 2). Tail long (longer than hind foot) and always clearly protrud- 

ing from the fiir. Body length less than 1.2 m. 
57 4 ( 5). Tail distinctly bicolored — 6-7 dark transverse rings distinct 

against lighter background 

Raccoon family, Procyonidae.* 

5(4). Tail one color, or terminal part darker than base, but trans- 
verse dark rings absent 

Marten family, Mustelidae.* 

6(1). Four digits on hind foot. 

7(8). Four digits on fore foot. Along neck and back, high, protrud- 
ing, shaggy crest of long elastic hairs. Color gray (without 
reddish brown tones), with black stripes across the trunk. . . . 
Hyaena family, Hyaenidae'^. 

8(7). Five digits on fore foot (first located somewhat higher than 
remainder). High crest along neck and back absent. Color 
different; if black transverse stripes on trunk, reddish brown 
tones well-developed. 

9 (10). Head elongated, with long dog- or fox-like muzzle. Tail 
always reaches tarsal joint; fluffy, but hairs short at base, 
thus tail is thinner at base than in middle. Claws not retrac- 
tile, their tips directed forward and downward, slightly 
compressed laterally and relatively blunt. Color without dis- 
tinct s^ots and transverse stripes 

Wolf family, Canidae (p. 66). 

♦Described in Part lb. 

'^Described in the following volume. 



65 



10 ( 9). Head round with short, cat-like muzzle. Tail reaches the 
tarsal joint, or is shorter, covered with hairs of equal length; 
at base, not thinner than in middle. Claws retractile, their tips 
directed forward and upward, strongly compressed laterally, 
and very sharp. If claws not retractile, then of large size 
(large dogs) and the color is light with small distinct, irregu- 
lar black spots. Color with spots or transverse stripes, or 
unicolor Cat family, Felidae.'^ 



Identification by Skull 



1 ( 6 



58 



2 ( 3 



Palatine bones protrude backward behind transverse line be- 
tween posterior points of last teeth, and less than half [palatine] 
width lies between these teeth'"* (Fig. 12). 
Four teeth in lower jaw on each side behind canine, five in 

upper; total number of teeth 34 

Hyaena family, Hyaenidae.*^ 

Dental formula different. 

Three teeth on each side in lower jaw behind canine; 4 or 3 

in upper; total number of teeth 28 or 30 

Cat family, Feiidae.'^ 

Seven or 6 teeth behind canine in lower jaw, 6 in upper; total 

number of teeth 42 or 40 Wolf family, Canidae (p. 66). 

Palatine bones protrude behind transverse line between pos- 
terior points of last teeth; more than half their width lies 
between these teeth "• (Fig. 12). 
Size of skull large. Maximum length more than 200 mm. . . . 

Bear family, Ursidae (p. 586). 

Size of skull smaller. Maximum length less than 200 mm. 
Six teeth behind canine in upper jaw and 6 in lower; total 

number of teeth 40 Raccoon family, Procyonidae.* 

10 ( 9). Less than 6 teeth in upper jaw behind canine; 4, 5 or 6 in 

lower; total number of teeth 38 or less 

Marten family, Mustelidae.* (V.H.). 



5 ( 4 

6 ( 1 



7 ( 8 

8 ( 9 

9 (10 



'^Described in the following volume. 

'"Measurement taken along suture between palatine bones. 

*Described in Part lb. 



66 





Fig. 12. Skull of fox, Vulpes vulpes L., and wolverine Gulo gulo L., on right 

(schematic). Characteristics of palatal construction and relation of the posterior 

part of tooth row and palate. Sketch by N.N. Kondakov. 

Superfamily of Wolves 
Superfamilia CANOIDEA Simpson, 1931 

Family of Wolves 
Familia CANIDAE Gray, 1821 

Carnivores of a more or less generalized type, of medium size, with 
unusually long le^s, light, well built and highly mobile. 

Digitigrade; tarsal and сафа1 regions never touch the ground dur- 
ing walking. On the anterior extremities there are five (in one case 4; 
Lycaon) digits, of which I is always rudimentary and located consid- 
erably higher than the others; on the posterior extremities, four digits. 
The two middle digits are considerably longer than the rest. The claws 
are not retractile, relatively blunt and not shaфly compressed laterally. 
The last phalanges are not capable of flexing far back and upwards and 
have normal articulating surface. 

The skull with long facial portion; small only in exceptional spe- 
cies; swollen auditory bulla, with a more or less horizontal internal 
partition, and the bone of the auditory meatus is of moderate length. 
The lateral occipital (paroccipital) process is small, somewhat laterally 



67 

compressed, and adheres to the auditory bulla. The mastoid process is 
small. The alisphenoid canal is well developed. 

The dental system is relatively complete* the usual dental formula 

is: I-cip-M- = 42. In one species it is I-cip-M- = 38 
3 14 3 3 14 2 

3 14 3-4 
(Speothos') and in another, it is: 1-C-P-M-— - = 46-48-50 
^ ^ 4 14 4-5 

{Otocyon). 

59 The camassial teeth are well developed, whereas the molars are 

short and quite flat with low blunt cusps; in the upper ones, their width 

exceeds their length. They have a grinding character, more or less 

"omnivorous". The canines are highly developed and are sometimes 

very long and thin. 

The caecum is fairly large "and spirally coiled or very weakly 
developed and hardly visible. The glans penis is long and smooth 
(without spines). An os penis is present, straight and bears a groove on 
its upper surface. 

The head is usually elongated with a long tapering muzzle. The 
ears usually have a shaф apex and are located fairly high, of moderate 
length, sometimes very long (longer than the head) and, as an excep- 
tion, they may be short with rounded apex. The tail is of moderate 
length (to tarsal joint) or longer, rarely short, and is covered all along 
its length with long hairs (the hairs are shorter at the base). Cutaneous 
glands are weakly developed, the anal glands are few, sometimes ab- 
sent, and there are no genital glands; in some species, there is a scent 
gland on the upper side of the tail ("violaceous" gland). A scrotum is 
present. There are 3^4—5 pairs of teats. 

The pelage of northern forms, in winter fur, is long, dense, fluffy 
and frequently silky, whereas in the south it is usually more coarse and 
sparse. The general color is usually more or less monotone, gray, 
yellow and red tones, sometimes with a dark dorsal saddle; and as an 
exception, almost black or of large and bright irregular black, white 
and red spots. In some species there are several color phases. 

There is no sexual dimorphism; age dimoфhism in some species is 
rare in some or is quite 8Ьаф1у distinct. Seasonal dimoфhism in forms 
of temperate and northern latitudes is considerable, manifested in the 



'According to some data, this species has 36-38-40 teeth (Haltenorth, 1958). 



68 

density and length of fur, and sometimes, in color (arctic fox). There 
are 1-2 molts per year. 

In general constitution, species of the family are homogeneous, 
representing a variation of one type as a whole — the wolf type. The 
change in appearance is mainly determined by the proportion of the 
trunk length and that of legs, and in part, tail length. 

The form with the most extremely long legs, resembling the borzoi 
dog, with a proportion of trunk length to shoulder height of about 1:1, 
is represented by the South American maned wolf {Chrysocyon 
brachyurus). The hunting dog of Africa {Lycaon pictus) is very close 
to it. The wolf is of the same type, but with shorter legs. The subse- 
quent type is represented by our fox and related species, which are of 
still shorter legs and elongated trunk, with proportion equal to 1 : 0.5; 
they have a long tail. The last group is represented by species which 
are similar to the raccoon dog {Nyctereutes procyonoides), with the 
extreme in this form represented by the South American bush dog 
(Speothos venaticus). It has a somewhat "dachshund-like" appearance, 
with proportions of 1 : 0.3 or slightly more, and a short tail. 

The species differ considerably in size. The largest species — ^wolf 
(Canis lupus) — has, in its large forms, body length to 1 60 cm, tail to 
50 cm, height at the shoulders to 85 (100) cm and weight to 50-60 kg 
or sometimes, exceptionally, to 80 kg. The smallest form — ^fennec 
(Fennecus zerda) — ^has a body length of about 40 cm, tail of about 20 
cm, height at the shoulders is about 20 cm and weight of about 2 kg. 
The proportions of extreme forms by weight is, in this way, about 
I : 25-30. 

The wolf family in its full breadth of specialization, both as a 
whole and also in particular species, represents one of the relatively 
"generalized" groups of the order. 

The majority of the species of the family are excellent runners, 
usually catching their live prey in open pursuit; some of them are very 
quick and have extremely great endurance in running. They are not 
60 adapted to climbing trees, and only the gray fox of America (Urocyon) 
can climb somewhat. Some dig quite well. They live in all landscapes, 
up to extreme high montane, but the majority, in all continents, are 
connected with the more or less open areas (forest steppe, steppe, 
desert). They frequently live individually, some live in pairs, and some 
species apart from the breeding season live and hunt in family groups 
or packs which are sometimes of great significance (hunting dog, Lycaon 
pictus; red wolf [dhole], Cuon alpinus). 



69 

The majority are sedentary, but some species accomplish consid- 
erable seasonal or irregular migrations or lead a nomadic life. They are 
monogamous animals, some species live in pairs for many years. The 
number of offspring [per litter] is up to 13—18 (according to some data 
up to 22-23), but is usually 4—7 and not more than 10. Males of the 
majority of species take part in the rearing of their offspring. This 
family is psychologically one of the most highly developed groups of 
carnivores, with definitely amicable internal relationships in packs and 
families, hierarchies, etc. This is, to a considerable degree, associated 
with the life in packs of many of the species. 

They are principally nocturnal, frequently crepuscular and rarely 
diurnal. Vision is relatively weak, hearing very good, and the sense of 
smell is shaф to the highest degree. The fresh meat of small and 
middle-sized mammals constitutes their main food, but several feed on 
large ungulates, carrion and rarely insects and other invertebrates. They 
utilize plant food, mainly fruits and berries, as supplementary food. 

The geographic distribution of the family is exceptionally vast — 
practically almost cosmopolitan. It corresponds to the geographic range 
of the order; however, the species of this family are absent from 
Madagascar, the entire Malayian archipelago with the exception of 
Sumatra and Java, the Philippines, Hainan, Taiwan and the Ryukyu 
islands. Occurrence in Australia (dingo) is secondary as a result of 
domestic dogs becoming feral (see characteristics of the order). In the 
north, the representatives of the family inhabit all lands and are found 
on oceanic ice nearly to the pole (arctic fox). On the Falkland Islands, 
they are extinct; several species (wolf) had their ranges reduced; how- 
ever, the geographic area of the family as a whole did not essentially 
change since historical time. 

The family represents an ecological quite plastic and quickly adapt- 
ing flourishing group of carnivores. The decline in numbers as well as 
range reduction of some species occurs only as a result of excessive 
hunting and destruction. Besides, the wolf, for example, persists even 
in places with special hunts for its total destruction. 

According to its systematic characteristics, the family occupies a 
quite distinct position and is shaфly distinguished from other families. 
It is considered to be closest to the bear family, Ursidae. However, 
judging by some characteristics of the genus Nyctereutes (raccoon 
dog), the relation of this family to that of raccoons (Procyonidae) may 
be closer (Frechkop, 1959). The resemblance of some forms to the 



70 

hyaena, Hyaenidae, is purely external, and is, to a certain extent, con- 
vergent. 

Concerning their origin, Canidae is one of the most ancient branches 
of carnivores, already identifiable in the upper Eocene. The earliest 
forms {Procynodictis, Cynodictis) are very primitive and possess fea- 
tures indicating their relation to forms which lie in the root of all 
Camivora = Fissipedia. 

Concerning the internal systematic structure of the family, it. is 
quite unified. Attempts were made to divide it into some families by 
separating such forms as the red wolf (Cwo«) and hunting dog (Lycaon), 
bush dog (Speothos) and, mainly, the bat-eared fox (Otocyon); how- 
ever, these attempts were poorly based and did not meet with acceptance. 

The present division into subfamilies is fairly simple and is actu- 
62 ally generally applicable. There are usually six subfamilies: the recent 
Caninae (wolves, foxes, raccoon dogs), Simocyoninae (Cuoninae, 
Lycaoninae — red wolves and others), Otocyoninae (bat-eared foxes) 
and three extinct ones: Amphicyoninae, Amphicyonodontinae and 
Borophaginae (Simpson, 1945). Sometimes, the genus Speothos (South 
American bush dog) which, in the above-mentioned system is related 
to Simocyoninae, is considered as a separate family — Speothoninae. 
Among the existing forms, this species is so peculiar that this point of 
view, supported most recently (Haltenorth, 1958), is represented as 
most justified. 

However, this system — ^at least that concerning the recent forms — 
is not irreproachable. If the justified separation of the genus Otocyon 
with its extremely peculiar dental formula in a separate group is ap- 
proved (see above), this cannot be applied in relation to Simocyoninae. 
The genus Lycaon (hunting dogs) of this group has the same dental 
formula as all other Caninae (42) but it has only four digits on the fore- 
limbs (the metacaфal bone of the first digit is even reduced); genus 
Cuon (red wolf) has the same number of digits on the forelimbs as in 
Caninae, but the dental formula is somewhat reduced (40, see below). 
The differences in the skull structure of the members of these genera 
from that of all Caninae in general, are not more than the differences 
between the various forms of Caninae. The differences in the structure 
of MI (see below) are also not very considerable and have no principal 
significance. According to what has been previously mentioned, there 
is no shaф limitation between Simocyoninae and Caninae in the char- 
acteristics mentioned. 



72 

The genus Speothos is better differentiated, first of all, by its 
dental formula (38, see above), which is still more reduced than that 
of Cuon, and by its "dachshund-like" appearance, blunt head, rounded 
ears, short tail, etc. This form is shaфly different, not only from Caninae 
and Simocyoninae {Cuon, Lycaon), but also from the genera with which 
it is frequently included in one group. Combining the characteristics in 
the family allows us to think that it is more natural to divide this family 
into two subfamilies — Otocyoninae (Otocyon) and Caninae (all other 
Canidae) — or to three, but in a combination of form different from the 
adopted one: with the retention of Cuon and Lycaon in the Caninae, but 
with the separation of Speothos in a special subfamily — Speothoninae.^ 

The number of genera in the family estimated by various authors 
is different. Lately, 68 genera are accepted, of which 56 are extinct and 
12 existing. Of the latter, eight genera belong to Caninae (Nyctereutes, 
Canis, Alopex, Vulpes, Fennecus, Urocyon, Dusicyon, Chrysocyon), 
three to Simocyoninae {Cuon, Lycaon, Speothos) and one to Otocyoninae 
{Otocyon) (Simpson, 1945). Some, mainly on account of the American 
forms, employ a greater number of genera. The actual number is, ap- 
parently, 7—8: Caninae — Nyctereutes, Canis, Chrysocyon; 
Simocyoninae — Speothos, Lycaon, Cuon; Otocyoninae — Otocyon. The 
other genera usually employed are only subgenera, mainly of the genus 
Canis (see below, Haltenorth, 1958).^ The total number of species in 
the family which was considered not long ago to be fairly large (about 
45), equals 29; namely, 23 species of Canis, Alopex and Vulpes — 
without the dingo and, excepting further reallocations in generic 
characteristics — simensis (subgenus Simenia), rueppelli, chama, 
pallida, zerda (subgenus Fennecus), microtis, thous (subgenus 
Lycalopex), cinereoargenteus, culpaeus, hagenbecki, australis 
{Dusicyon), Chrysocyon — 1 (C. brachyurus — South American maned 
63 wolf), Nyctereutes — 1 (see below), Speothos — 1 {S. venaticus — South 
American bush dog), Lycaon — 1 {L. p ictus — ^African hunting dog), 
Cuon — 1 (red wolves; see below), Otocyon — 1 {O. megalotis — ^Afri- 
can bat-eared fox). 



^Systematic relationships in the family require special analysis, and here the 
traditional grouping of the species of our fauna into two subfamilies is conditionally 
given. 

^The uniting of the genera Cuon and Lycaon. even without the application of 
subgeneric rank, suggested by the same author, is not accepted. It is possible that 
their union was based on the fact that the author wrongly applies the same dental 
formula for both genera (42), although in Cuon there are only 40 teeth. 



73 

Six species inhabit Asia {V. сапа, V.ferrilata, V. bengalensis* , V. 
corsac, N. procyonoides, C. alpinus); in Africa — ^seven (C. simensis, 
K** chama, K** pallidus, C. adustus, C. mesomelas, L. pictus, O. 
megalotis); in South America — seven {D.^^*^ microtis, D.*** thous, D. 
culpaeus, D. hagenbecki, D. australis, С brachyurus, S. venaticus); 
in North America — two (V. velox, C. latrans). One species lives in 
Eurasia, Africa and North America {V. vulpes), two species in Africa 
and Asia (V** nieppelli, F.** zerda), two in Eurasia and North America 
(C. lupus, A. lagopus), one — ^in Africa and Eurasia (C aureus) and 
one- — ^in North and South America {D. cinereoargenteus). 

Among the species of the family there are exploited animals which 
give fur, usually expensive, and — ^to a lesser degree — ^are objects for 
sport hunting. Some species are important destroyers of rodents. A 
considerable number of them have no noticeable utility and, in general, 
any practical significance. Some species are dangerous pests for live- 
stock breeding and the hunting economy, and some are reservoirs for 
rabies virus, transmitting it to domestic animals and man. 

One of the species (wolf) has its geographic range considerably 
decreased in connection with its eradication in many places both in the 
Old and New World, and another one, Falkland wolf, Canis (Dusicyon) 
australis, is completely extinct. The geographic range of some species 
has increased in connection with their acclimatization (red fox). One 
species is present in a wild form as well as in a completely domestic 
form (wolf-dog) and two species are present in a wild and semi-domes- 
tic form as objects of fur-farming (red fox and polar fox). 

In the USSR fauna, 2 of the 3 subfamilies are represented: Caninae 
and Simocyoninae; 5 of the 12 existing genera (42%)'*: Nyctereutes, 
Canis, Alopex, Vulpes, Cuon, and 8 species of 29 (about 27% of the 
species of the family); they constitute about 2.6% of the total number 
of species of our fauna. 

The geographic range of this family covers the whole territory of 
the USSR and its species are found under all natural conditions, to 
extreme high montane regions and the ice of the Arctic Ocean. 



*Misspelled lengalensis in Russian original — Sci. Ed. 

** Spelled F. 'probably for subgenus Fennecus, in Russian original — Sci. Ed. 

***Spelled L, probably for subgenus Lycalopex, in Russian original — Sci. Ed. 

The treatment of genera and subgenera is inconsistent — Sci. Ed. 

"Or 3 out of 7, i.e., 43% 



74 

The species of the family have an important value in the USSR as 
fur-bearing and sport hunting animals, destroyers of rodents, pests for 
animal breeding and hunting and carriers of some diseases (V.H.). 

Key for Species Identification of the Wolf Family 

1 (2). Six teeth (premolars and molars) behind canine on lower jaw; 
total number of teeth 40. Condylobasal length of skull not 
less than 170 mm. General dimensions slightly less than those 
of wolf. Color bright reddish-yellow; dorsal side of ear of 
same color. On fore limbs, pads of middle digits connected 

posteriorly by an anastomosis (Fig. 17) 

red wolf [dhole], Cuon alpinus (p. 571). 

2(1). Seven teeth behind canine in lower jaw; total number of teeth 
42.^ Color different; if red, dorsal side of ear black. 

3 ( 4). Posterior lower angle of lower jaw below angular process 

with sharp, deep angular groove (Fig. 14). Muzzle with dark, 
mask-like shading, also covering eye region. Size similar to 
fox raccoon dog, Nyctereutes procyonoides (p. 82). 

4 ( 5). No shaф angular groove on posterior lower angle of lower 
64 jaw, or only a very gently sloping, not deep arciform groove. 

Dark "mask" on muzzle covering eye region absent. 

5 ( 8). Frontal (interorbital) region of skull noticeably elevated rela- 

tive to nasal region, and line of upper profile of skull forms 
protrusion here. Postorbital processes convex above. Apex of 
lower canine, when jaws are closed, does not reach margin 
of alveolus of upper canine. Facial part of skull relatively 
short and broad (distance from posterior margin of preorbital 
foramen to posterior margin of canine alveolus less than width 
of skull above canines). 

Tail with hair equal to or shorter than half of body length. 
On undersurface of hind foot, five large bare pads (callosi- 
ties), four digital and one behind them. Pupil rounded. 

6 ( 7). Arcuate line formed by anterior margins of nasal bones lacks 

sloping middle process (near suture of both bones). Protrud- 
ing cingulum on outer side of base of first upper molar (second 
tooth from rear) narrow and blunt with median constriction or 



'As an individual difference, the number of molars and premolars may be 6. 
However, in this case, the longest skull is 125 mm. 



75 




Fig. 14. Lower jaw of fox, Vulpes vulpes L., and raccoon dog, Nyctereutes 

procyonoides Gray, below (schematic). The deep notch on the lower posterior 

angle of the jaw of raccoon dogs is shown. Sketch by N.N. Kondakov. 

gap. On fore limbs, pads of two middle digits not united by 
commissure in posterior part (Figs. 15, 16, 17). 

Dimensions large, skull length more than 200 mm; total 
body length of adults more than 90 cm. Tail length almost 

half of body length wolf, Canis lupus (p. 164). 

7 ( 6). Arcuate line formed by anterior margins of nasal bones has 
in middle (at suture of bones) small blunt and anterior pro- 
jection. Cingulum on base of first upper molar wide, shaфly 
defined and without gaps. On fore limbs, pads of two middle 
digits are connected by commissure in posterior part (Figs. 
15, 16, 17). 

Dimensions smaller, skull length not more than 190 mm; 
total body length of adults less than 90 cm.^ Tail length is 
about half that of body .... jackal, Canis aureus (p. 129). 
65 8 ( 5). Frontal (interorbital) region of skull not noticeably elevated 
relative to nasal region, and line of upper profile of skull 
relatively uniformly elevated towards rear. Postorbital pro- 
cesses flat or concave above. Apex of lower canine, when 



*In the species, as a rare exception, body length may be somewhat larger. In such 
case, identification is decided on skull size, structural craniological characters, and 
structure of the pads of the fore limbs. 



76 





64 Fig. 15. Anterior margin of nasal bones of wolf (to the left) and jackal — dorsal 
view. Sketch by N.N. Kondakov. 




65 Fig. 16. Cingulum of anterior upper molar of wolf (above) and jackal. Sketch by 

N.N. Kondakov. 





65 Fig. 1 7. Sole and digital pads of fore limb of wolf, jackal and red wolf (left 
to right). Sketch by N.N. Kondakov. 



77 

jaws are closed, reaches margin of alveolus of upper canine 
or extends beyond. Facial part of skull relatively narrow and 
long (distance from posterior margin of preorbital foramen to 
posterior margin of canine alveolus more than or equal to 
width of skull above canines). 

Tail with hair exceeds half of body length. Undersurface 
of hind foot has only 4 circular, bare digital pads (callosi- 
ties). Pupil oval, vertical. 

9 (10). Postorbital processes above flat or slightly concave. Apex of 

lower canine, when jaws are closed, reaches only margin of 
alveolus of upper canine or extends a bit farther above it 
(Fig. 18). Distance from posterior margin of infraorbital 
foramen to posterior margin of canine alveolus equal or 
almost equal to skull width above canines. 

Ear with rounded tip, relatively short; when bent for- 
ward, tip does not reach eye. Body color either completely 
white or evenly brownish-gray (winter fur), or back, limbs 
and tail are brownish while side and venter are dirty white 
arctic fox, Alopex lagopus (p. 274). 

10 ( 9). Postorbital processes above strongly concave (at base, dorsal 

groove is formed). Apex of lower canine, when jaws are 
closed, extends considerably past margin of upper canine 
alveolus. Distance from posterior margin of infraorbital fora- 
men to posterior margin of canine alveolus greater than width 
of skull above canines. 

Ear with shaф tip, long; when bent forward, tip reaches 
eye. Color different. 

11 (12). Posterior side of ear brownish-black or pure black and shaфly 

distinguished from red or sandy-reddish color of back. Black 

markings on anterior side of fore limbs. Extreme tip of tail, 

66 at least very small part, white. Size relatively large; body 

length more than 65 cm, total length of skull more than 1 1 5 

cm, width of zygomatic arches more than 64 mm 

red fox, Vulpes vulpes (p. 469). 

12 (11). Posterior side of ear not black or brownish-black and not 

distinguished from color of back. No black marks on anterior 
side of fore limbs. Tail tip not white. Size smaller. 

13 (14). Back gray mixed with black (in form of ripples). Chin and 

lower lip dark brown or black. Canines very long; when jaws 
are closed, ends of upper canines protrude beyond level of 



78 






Fig. 18. Canines and incisors from side* (Sketch by N.N. Kondakov). 1 — ^wolf; 

2 — jackal; 3 — arctic fox; 4 — Middle Russian red fox (Vulpes vulpes vulpes L.); 

5 — ^Turkmenian red fox (Г. v.flavescens Gray); 6 — Corsac fox. 

lower margin of lower jaw. Size very small; body length less 
than 50 cm, skull length less than 95 mm, zygomatic width 

less than 50 mm Afghan fox, Vulpes сапа (p. 390). 

14 (13). Color of back is rusty or rusty-yellowish, grizzled from the 
white ends of the guard hair. No mixture of black color. Chin 
and lower lip white. Canines relatively shorter. Ends of upper 
canines, when jaws are closed, reach level of lower margin 
of lower jaw. Size larger; body length more than 50 cm, skull 
length more than 95 mm, zygomatic width more than 58 mm 
corsac fox, Vulpes corsac (p. 396) (V.H.). 

Subfamily of Wolves 
Subfamilia Caninae Gill, 1872 

Genus of Raccoon Dogs, or Manguts 
Genus Nyctereutes Temminck, 1839 

1839. Nyctereutes Temmink. In: Van der Hoevens Tijdschr. Nat. Ges. 
Phys., 5, p. 285. Nyctereutes viverrinus Temminck. 

Forms of small size. 

Skull relatively quite heavy and massive, moderately elongated, 
with narrow zygomatic** arches, and not high (height in occipital 



*ln Russian original, "front", an apparent lapsus — Sci. Ed. 
**"auditory" in Russian original, an obvious lapsus — Sci. Ed. 



79 

region little more than one third of condylobasal length). Protuberances 
and crests of skull well developed, sagittal crest high in old animals and 
occipital quite considerable. 

Facial region of skull short and its length nearly equal to cranial — 
distance between anterior end of premaxillae and line connecting ends 
67 of supraorbital processes equal or nearly equal to that between this line 
and upper margin of foramen magnum. Facial region of skull massive. 
Distance between posterior wall of infraorbital foramen and posterior 
margin of canine alveolus less than width of skull above canines or 
equal to it. 

Interorbital (frontal) region elevated, somewhat swollen in region 
of frontal sinuses between supraorbital processes; processes themselves 
not large, flat or slightly concave above, with tip sharply recurved 
downwards. Upper profile of skull at first almost straight and then 
becomes convex arcuate line; at anterior of nasal region already abruptly 
elevated and no frontal shelves occur. Posterior edge of tooth row does 
not reach level of anterior border of inteфterygoid notch. Hard palate 
extends backwards behind line uniting posterior points of last molars 
to approximately 1/3 the distance between inner comers of last molars. 

At lower edge of lower jaw, anterior to angular process, a large, 
massive protuberance, separated from angular process by deep angular 
notch. Such a mandibular structure also present only in American gray 
fox, Dusicyon (Urocyon) cinereoargenteus* (Fig. 14). Angular pro- 
cess large and massive, with broad base and without upwardly directed 
denticle. 

3 14 2 
Dental formula: I— C-P— M— = 42. As a fairly rare individual 

3 14 3 

3 
deviation, there may occur M— = 44. Teeth relatively small and weak. 

3 

Upper canines strong, but short, their ends far short of upper border of 
lower jaw, and slightly curved; their ends not directed straight down- 
wards, but somewhat slanted to the side. Cutting edge on posterior side 
of canine and on lateral surfaces, a blunt, smooth vertical edge. Upper 
camassial teeth relatively weak. Incisors have no additional blades; 
external [third] incisor not adjacent to second, and separated from it by 
noticeable gap. 

*Mispelled cinereoargentatus in Russian original- — Sci. Ed. 



80 

Trunk long, legs short, and tail hangs below tarsal joint, but when 
the animal is standing, does not reach ground. Ears of moderate length. 

Pelage (in northern forms) dense and fluffy in winter. Molt occurs 
once a year. Five pairs of teats. Intestine relatively long, compared to 
body length being 4.3 : 1 (Komeev, 1954). 

Inhabitants of closed landscapes — ^forests, river valleys, preferably 
moist. Of low mobility, and travel relatively slowly. They do not per- 
form regular migrations. Monogamous animals, pairing for one breeding 
season. 10-15 offspring, usually less. Young bom in burrows, and they 
are associated with the burrows throughout the entire year. Apart from 
the period of reproduction, they live singly. They are predators, feeding 
on small vertebrates, especially rodents, amphibia, eggs, insects and 
other invertebrates, and also on berries and fruits. In the northern part 
of their range, they hibernate (the only species in the family). 

The natural range of this genus is not great and is limited to part 
of eastern Asia. It occupies the southern parts of the Amur [and] Ussuri 
territories, northeastern China (former Manchuria), the Korean penin- 
sula, eastern China to the extreme south, and the Japanese islands. 

In the last ten years, the range has significantly expanded due to 
acclimatization in almost the whole European part of the [Soviet] Union 
and in the Caucasus, in separate places in southern Siberia and the 
eastern part of Middle Asia (see below; description of species). 

Because of its systematic characteristics the genus Nyctereutes 
represents a strongly deviant form of the subfamily and stands com- 
68 pletely apart. In any case the genus is more isolated from other genera 
of the subfamily than they are from each other. Even by broad inter- 
pretation of the genus, when such groups as Vulpes, Alopex, Fennecus, 
Simenia, Lycalopex and Dusicyon are included in the genus Canis (see 
characteristics of the family), the generic independence of Nyctereutes 
remains obvious and undoubted. 

The genus may be considered specialized in some respects; how- 
ever the more basic features, mainly in the structure of the facial part 
of the skull, in dentition, and in some other characters, are primitive. 
The genus ought to be included among the least specialized in the 
subfamily (family ?) and at the base of the systematic series. Also 
primitive are such characteristics as poorly developed predatoriness, 
low mobility (the animal is essentially not a predator, but a "collec- 
tor"), large number of young and hibernation. Apparently, it is 
sufficiently basic to consider the genus as archaic and relict. This is 
supported by its geographic range, which is associated with a region in 
which Tertiary relict forms are preserved. 




Fig. 19. Geographic range of the mangut, or raccoon dog , Nyctereutes Temm. 
In the east, the natural species distribution of the mangut; in the west, the region 
inhabited by acclimatized animals in the European part of the Soviet Union and 
in the Caucasus. Arrows point to the direction of spread of the raccoon dog from 
the USSR to Finland, Poland, Romania and Central Europe. (V.G. Heptner). 

69 Nyctereutes has some characters similar to those of the raccoon 
family, Procyonidae. Thus, both in general outlines and some structural 
details, the skull of Nyctereutes resembles that of Bassariscus 
(cacomistles), and the dentition occupies, in a certain sense, an inter- 
mediate position between that of the true Canidae (Vulpes) and 
Procyonidae (Procyon). The biological characters noted, especially 



82 

hibernation, are also "raccoon" features (Frechkop, 1959). These char- 
acters do not provide a basis for including Nyctereutes in the raccoon 
family, but they well emphasize the individuality of the genus. More- 
over, they show that the relationship between the two named families 
is closer than what is usually believed. 

Fossil representatives of the genus are not known. However, it can 
be assumed that the time of their development is concerned with the 
Pliocene, and perhaps even earlier. 

The genus included one species: Nyctereutes procyonoides (Gray, 
1834),* i.e. about 3.4% of the species of the family. This species 
constitutes about 0.3% of the species of the USSR fauna. 

The natural range of the genus in the USSR is restricted to a small 
strip of forests of the Manchurian type in the Ussuri and Amur terri- 
tories. This species has been acclimatized in other parts of the USSR 
(see below). 

It is one of the secondary fur-bearing animals; usefiil to a certain 
degree in destroying rodents, but it is essentially harmful to the wild 
game economy (V.H.). 

USSURI RACCOON, RACCOON DOG, MANGUT^ 

Nyctereutes procyonoides Gray, 1843 

1843. Canis procyonoides Gray. Illustr. Zool., 2, pi. 1. Vic. Canton. 
1907. Nyctereutes ussuriensis. Matschie, Wiss. Ergebn. Filchners. Exp. 

nach China, 10, p. 178, Near Ussuri mouth. 
1907, Nyctereutes amurensis. Matschie. Ibid., p. 179. Amur. 
1958. Nycterepites procionoides kalininensis. Sorokin. Byull. Mosk. 

obshch.'ispyt. prirody. Kalinin Department. 1. Kalinin district. 

(V.H.). 

Diagnosis 

Only species of the genus. 

*In Russian original "Temmink, 1839" — Sci. Ed. 

^First and second names are artificial and literal. "Raccoon dog" is simply a 
translation from the Latin — Canis procyonoides. It is the most common name in our 
literature. "Ussurian raccoon" is a ftirrier's term. The Russian local (Amur Cossack) 
name is "enot" [raccoon] (Cherkasov, 1867). Mangut is an Evenk name. It is better than 
the others, and was used by Russians. Unfortunately, it is not used in literature at the 
present time. 



83 

Description 

General appearance of the Ussuri raccoon is entirely unique, not resem- 
bling the appearance of other representatives of the family. The trunk 
is long, legs short and they look particularly short in the animals with 
winter fiir, which on the trunk is very long. Accordingly, the body has 
a massive appearance. Also, the legs are covered with short hairs and 
they look very thin. This gives the impression of a stocky and clumsy 
animal on short, thin legs. 

The fur in winter is not proportional to the size of the animal. It 
is long (length of guard hair to 120 mm), thick, with dense underfur and 
coarse guard hair, shaggy and stiff. The tail is relatively short — not 
more than 1/3 the body length, its end falls below the tarsal joint, but 
it does not reach the ground when the animal is standing. The hairs of 
the tail are very long and dense, not shorter at the base than on any 
70 other part of its whole length and the tail fiir is not clearly distinct from, 
and is transitional to, the fur of the croup. This reinforces the impres- 
sion of a general elongation of the animal's body. The head is not large, 
with short, but pointed muzzle covered with short hairs. Behind the 
eyes the hair rapidly increases in length and quantity, and quite large 
ears, blunted on the ends, protrude slightly from the fiir. There are large 




Fig. 20. Mangut, or raccoon dog, Nyctereutes procyonoides Gray. Sketch by 

A.N. Komarov. 



84 



■ '^^S^'^^sf- ¥'.¥^ 



-I 



Ш 




Fig. 21. Male mangut, 10-11 months of age. "Kedrovaya Pad' " preserve 
(southern Primor'e). March 1964. Photograph by A.G. Pankrat'ev. 

"side-whiskers" of elongated hairs on the cheeks. The pointed muzzle 
looks as if it is emerging from the long fur of the head. 

The general color tone of the winter fur is dirty, earth-brown or 
brownish-gray with a more or less considerable overlay of black (color 
of guard hairs). The tail is considerably darker than the trunk. Along 
the back extends a darker stripe which broadens on the shoulders, 
forming there an unclear cross-shaped figure. The abdominal surface is 
yellowish-brown and the chest is dark brown or blackish. Ears are 
black posteriorly. In the eye region, in front of them and on the cheeks 
below and behind the eyes and crossing the "side-whiskers", is located 
a dark (almost black) field. Together, they form on the muzzle of the 
mangut a characteristic picture in the form of a mask which contrasts 
8Ьаф1у with the brighter color of the muzzle and remaining parts of the 
head. This picture in particular, and the general color tone gives the 
described species a certain similarity to the American raccoon. 
72 In summer, during the transition from one winter coat to another, 

the mangut has at first (first half of summer) only a pelage of dry 
lifeless top hairs (underfur sheds first) and later, the fiir is formed of 
short, incompletely grown guard hairs, almost without underfur. Such 
hairs are several times shorter than winter ones. The color during that 



85 

time resembles that in winter, but its general tone is brighter, reddish- 
straw, but with a mixture of blackish-brown. The dark color forms a 
stripe along the back and dorsal side of neck, and a cross-shaped figure 
in the shoulder region. The chest is brownish-black while the belly and 
groin are brownish-gray with yellowish. The legs are blackish-brown, 
the "mask" is well defined; the chin, neck and chest are black. On the 
sides of the neck the color is quite bright, pale-rusty. Claws are light- 
horn. 

For description of skull, see description of the genus. The diploid 
number of chromosomes is 42.* 

Relatively speaking, data on the dimensions and weight of manguts 
are extremely scanty. Body length is about 65—80 cm; tail length, 1 5— 
25 cm; weight in summer 4-6 kg and in winter, 6-10 kg (C. Naumov 
and Lavrov, 1948). 

Dimensions of the skull (in mm) of males from the Far East are as 
follows (42 specimens; Sorokin, 1958): 

Range Mean 



Greatest length 


114.3- 


-131.8 


123.4 ± 


0.6 


Condylobasal length 


103.2- 


-125.8 


119.2 ± 


0.6 


Zygomatic width 


61.0- 


- 75.0 


67.2 ± 


0.4 


Width of muzzle between 


20.3- 


- 24.9 


22.3 ± 


0.1 


canines 










Width between orbits 


19.8- 


- 25.2 


22.6 ± 


0.1 


Distance between supraorbital 










processes 


26.0- 


- 36.0 


31.5 ± 


0.3 


Greatest width of skull 


40.4- 


- 44.5 


42.3 ± 


0.1 


Length of nasal bones 


35.4- 


- 50.7 


42.6 ± 


0.4 


Height of skull in region of 










auditory bullae 


41.3- 


- 48.3 


44.6 + 


0.2 


Length of upper tooth row 


42.7- 


- 49.4 


46.4 ± 


0.2 



Dimensions of the skull of females from the Far East 
(40 specimens): 

Range Mean 



Greatest length 


112.8-131.7 


120.3 ± 0.6 


Condylobasal length (36) 


110.1-126.6 


109.8 ± 0.5 



*Now known to be 38 + 1^ small В chromosomes for Japanese animals. Main- 
land raccon dogs have a diploid number of 54 + 1-3B chromosomes — Sci. Ed. 



86 




71 Fig. 22. Skull of mangut, or raccoon dog, Nyctereutes procyonoides Gray. 



Range 


Mean 




60.2- 71.3 


66.2 ± 


0.4 


18.7- 23.8 


21.5 ± 


0.1 


19.5- 25.5 


22.3 ± 


0.2 


38.9- 45.3 


42.1 ± 


0.2 


37.2- 49.7 


43.6 ± 


0.4 


41.0- 47.6 


44.6 ± 


0.2 


43.2- 51.7 


45.8 + 


0.2 



87 



Zygomatic width 

Width of muzzle between canines 
Width between orbits 
Greatest width of skull 
Length of nasal bones (39) 
Height of skull in region of 

auditory bullae (39) 
Length of upper tooth row 

Difference in the dimensions of males and females is so small that 
sexual dimorphism is basically not evident (V.H.). 

Systematic Position 

See characteristics of the genus. 

Geograptiic Distribution 

The forest areas of eastern Asia from the Amur territory to northern 
Indochina. This is a typical range for a species of the Manchurian- 
Chinese fauna. 

As a result of acclimatization, a new range was established in the 
European part of the [Soviet] Union and in the Caucasus, and there are 
separate sections of occurrence in Siberia and Middle Asia. 

Geographic Range in the Soviet Union 

The range in the USSR represents the extreme northern border of the 
range of the species. It is not large, and occupies a very small part of 
the country; namely the Ussuri territory and the southern part of the 
Amur. 

The outlines of the range within the territory of the [Soviet] Union 
are fairly complicated. In the east it begins at the Pacific coast of the 
Ussuri territory in the region of the mouth of the Tumnin river (about 
51° N. Lat.) and extends as a narrow strip along the seacoast to the 
73 south and southeast along the foothills and lower mountain belt. Pass- 
ing around the higher parts of the southern extremity of Sikhote-Alin' 
from the south and approximately in the latitude of the northern end of 
Lake Khanka, the boundary traverses the western slope of the range. 
Along the Ussuri valley and along the western slopes of Sikhote-Alin', 



the range strip passes north to the Amur and reaches it at the mouth of 
the Ussuri, Along the valleys of the more significant right tributaries 
of the Ussuri (Bikin, Khor, Iman), the range at places extends far to 
the east into the mountains; however, the mangut is absent in the 
extreme high-montane Sikhote-Alin', and in the region of the extensive 
coni-ferous forests. 

Along the Amur valley and adjacent places, the range extends 
northward (downstream) nearly to the Khungara. From this place the 
northern border of the range continues westward through the Burein 
mountains and the middle course of the Bureya and Zeya to the Amur 
in the region of the junction of the Shilka with the Argun. Because of 
scanty information, it is impossible to determine whether the species 
described is distributed all along the Amur valley to the west from the 
Zeya, or whether they penetrate from the south (from China) to the 
mouths of the Shilka and Argun. They are not found in the extreme 
northern part of the northern bend of Amur. According to some data, 
the northern border of the normal occurrence extends in our country 
from the east only to the mouth of the Khumaerkhe on the Amur. To 
the south, the range extends everywhere up to the state boundary. 

The mangut is absent in Trans-Baikal, although in remote times, it 
apparently reached the upper Onan.^ Sakhalin is not included in the 
natural range. 

In the last decades (starting from 1934), the Ussuri raccoon in- 
creasingly has been established, especially in the European part of the 
[Soviet] Union, and also transplanted to separate places in the Asiatic 
part. At the present time in the European part of the country, the 
mangut formed a vast distribution area, which considerably exceeds its 
natural range within the USSR. 

In the north the new range occupies all of Karelia and, probably, 
the entire southern part of the Kola peninsula, the whole of Arkhangel'sk 
district including the forest-tundra; however, incursions by the mangut 
onto the tundra occur. Apparently, the northern border passes some- 
what more to the south along the Pechora. In the west, the range in the 
entire European part of the Union reaches the western state boundary 
and in some places beyond it. In the south, the range reaches the Black, 
Azov and Caspian seas. The eastern border is less defined. On the 

^Old data about the permanent habitation of the animal here were not confirmed. 
The range is according to Cherkasov, 1867; Radde, 1862; Maak, 1859; Schrenk, 1858; 
Ognev, 1931; С Naumov and Lavrov, 1941; material of N.N. Rukovskii and other 
sources. 



89 

northeast, the mangut is known (settled) at the river Ishma (left tribu- 
tary of the Pechora). To the south, the border extends, apparently, 
along the Urals west of the range — through Perm district and Bashkiriya 
to Orenburg. From the Urals it extends to the Volga and follows it 
down to the [Caspian] Sea. The line of the eastern border of the 
European range is very undefined, not only due to insufficient informa- 
tion, but also because it changes as a result of introductions of the 
animal (easternmost transplantations: Syktyvkar, Udmurtiya, Ufa, 
Orenburg). 

Within the outlined borders, by 1960 mangut actually occurred 
everywhere. If it was still absent in more or less considerable parts 
(some parts of Karelia and Gis-Urals) or its distribution was sporadic 
or it was very rare (Arkhangel'sk district), then in most of the area 
(several districts) its number increased immensely and it became an 
important enemy of the hunting economy. 

In the Caucasus, the Ussuri raccoon has been transplanted 
and established in a series of places. Its range extends into the Cis- 
Caucasian steppes and foothill districts, and the eastern half 
of Trans-Caucasus. It is, apparently, absent only in the western part 
of Georgia. Its occurrence within these outlined boundaries is not 
continuous. 
75 Outside the above-described borders, Ussuri raccoons were estab- 
lished in Middle Asia and Kazakhstan — to the east of Frunze, in 
Issyk-Kul, in the Dzhelalabad region, at the mouths of the Hi and 
Lepsa, to the south of Alma-Ata, in the Dzhungarsk Alatau, and in 
Katon-Karaga region in the southwest Altai. 

In Siberia, manguts have been established at the mouths of the 
Biya and Katun, and in a series of places in the Altai, near Tom'sk at 
the sources of the Tom', near Novosibirsk, near Krasnoyarsk, in Tuva, 
at Irkutsk, northeast of Ulan-Uda and along the Dzhida and even in 
Yakutiya — ^at the mouth of the Vilyui and southeast of Suntar on the 
Vilyui. They were also acclimatized in several other places, including 
Sakhalin.^ Establishment in Siberia and Middle Asia did not lead to the 
formation of significant patches of occurrence, to say nothing of a 
continuous range. In the majority of places, apparently, the animals 
died or are dying, and in some places, small foci were formed. 

'Acclimatization of the mangut from data of Lavrov, 1946; Rukovskii, 1950; 
Komeev, 1954; Vereshchagin, 1959; Berens and Yakushevich, 1959; Marvin, 
1959; and a number of other works as well as the materials provided by N.N. 
Rukovskii. 



91 

Geographic Range outside the Soviet Union 

The range outside the USSR includes China, from former Manchuria 
southward to Fukien, Guandong and Yunnan, the Korean peninsula, the 
northeastern part of the Indochina peninsula (Tonkin, now Bakbo prov- 
ince in North Vietnam). Borders of the distribution to the west are 
undefined; in northeast China the range does not go beyond Great 
Khingan (absent in the Mongolian Republic); to the south, it is known 
from Shaanxi and Sichuan; absent in Burma. It is found in the Japanese 
islands (Honshu, Shikoku, Kyushu, but is absent on the Ryukyus). It is 
not present on Taiwan and Hainan. 

As a result of the spread of the animals acclimatized in the Euro- 
pean part of our country, at the end of the 1950's the Ussuri raccoon 
penetrated to Poland, Romania, Finland, northeast Sweden and even in 
the G[erman] D[emocratic] R[epublic], where they are found east of 
Berlin (V.H.). 

Geographic Variation 

Geographic variation of the species is not well studied. Five subspecies 
are usually accepted. Within the Soviet Union one subspecies is present. 

Ussuri mangut, N. p. ussuriensis Matschie, 1907 (syn. amurensis, 
kalininensis). 

For description, see above. 

Ussuri and Amur territories. Acclimatized in the European part of 
the USSR and in the Caucasus. 

Outside the USSR, northeast China and Korean peninsula; the 
limits of distribution to the south are unclear. 

This form is poorly described, but by comparison with the nominal 
form from the extreme south of the range it apparently is distinguished 
by the somewhat larger size and denser, longer hair. There is no com- 
parative study of our form. The name amurensis is based on an 
occurrence character. 



74 Fig. 23. Range of the mangut, Nyctereutes procyonoides Gray in USSR. 
Continuous line in the Far East is the northern border of the natural range of the 
species. The more or less continuous distribution formed as a result of 
acclimatization in the European part of the country and in the Caucasus is shown 
by the closed line in the West. Individual dots are the places of establishment of 
the animals beyond the limits of the western range. Data of the western part of 
the distribution area are from the beginning of the 60's. (V.G. Heptner). 



92 

Separation of animals from Kalinin district in the form of an inde- 
pendent subspecies N. p. kalininensis (Sorokin, 1959) was unnecessary 
and uncalled for. As is evident from the figures given by the author, 
there is no difference in the dimensions of the skull. Differences in the 
quality of fur (longer, fluffier, softer and denser) are very difficult to 
perceive. If differences exist, they mainly reveal completely normal 
phenotypic variability. The independence of this form has neither a 
practical nor theoretical basis. 

Outside the boundaries of the USSR, the following forms are usu- 
ally accepted: (1) N. p. procyonoides Gray, 1834 — China, except 
Yunnan, Tonkin; (2) N. p. viverrinus Temm., 1844 — ^Japanese islands; 
(3) N. p. koreensis Mori, 1922 — Korean peninsula; (4) N. p. orestes 
Thos., 1923 — ^Yunnan. The reality of the last two forms, especially the 
Korean one, is highly doubtful. Most probably there are 3 forms — the 
mainland southern {procyonoides), northern {ussuriensis) and island 
{viverrinus). (V.H.). 

76 Biology 

Population. There are hardly any data about the number of raccoon 
dogs within the limits of its natural range in the Far East. The densest 
population is observed in the southwestern part of Primorsk territory, 
south of Lake Khanka, in the lower and middle courses of the Iman, 
Bikin, and Khor rivers and the middle course of the Amur River. In the 
basins of the Maikhe and Suchan rivers and along the coast of the 
Japanese Sea south of Nakhodka Bay (Ussuri territory) in 1931, the 
population density (calculated by the number of burrows found) con- 
sisted in various habitats of from 0.9 to 8 animals, averaging 3.4 per 
1000 hectares. On Askol'd Island, density averaged 3.8 animals over 
the same area [i.e., 1000 h] (Bannikov, 1964). 

In the majority of regions of acclimatization of this species in the 
European part of the USSR, in the zone of mixed and deciduous forests, 
the population density constitutes about 1—2 animals per 1000 hectares. 
In the flooded deciduous forests of some regions, for example Novgorod 
(Morozov, 1953), it reaches 13 individuals per 1000 hectares; in the 
middle Volga region and Tatariya, 2—3 animals (V. Popov, 1953), in 
Gor'kovsk district, in different years, from 15 to 48 (Kozlov, 1952); 
in Voronezh district an average of 1—2 animals per 1000 hectares, but 
in some regions, for example Bogucharsk, to 20 individuals in such an 
area (Obtemperanskii, 1953; Ivanova, 1962). In the northeastern 



93 

regions of Lithuania, 5 to 10 animals were counted in 1000 hectares of 
forest area; in the northern regions, 2—4 individuals; and in the southern 
and western regions, about 0.5 per 1000 hectares (Prusaite, 1961). 

In Rostovsk district and in the Ukraine, the population density of 
raccoon dogs consists of 2— 5 animals per 1000 hectares of land. How- 
ever, on the forested islands and along the shores of water bodies, it 
reaches 29-^6 animals (Rail' and Kritskaya, 1953; Komeev, 1954; 
Sergeenko, 1956 and others). In the northern Caucasus, in Dagestan, 
Georgia and Armenia, Azerbaijan and Moldavia, the number of rac- 
coon dogs, as a rule, does not exceed 1—2 individuals per 1000 hectares, 
and frequently is much less (Stepanov, 1939; DaF, 1941; Meladze, 
1947; Kostyuchenko, 1950; Lavrov, 1957; Trushchalova, 1959; Aliev, 
1962 and others). In Kazakhstan and Kirghizia, the density is even 
lower but in small sections, for example, along the shore of Lake Issyk- 
КиГ, the raccoon dog population reaches 20-25 animals per 1000 
hectares (Berens and Yanushevich, 1956 and others). 

Habitat. In the Far East, preferred habitat of the raccoon dog is 
light deciduous and mixed forests near streams with dense understory, 
or the thick growths of shrubs, usually found on gentle slopes of moun- 
tains, interrupted by waterfalls and creeks, with rock outcrops and clear 
areas. In southern Ussuri territory, more than half of all encounters 
with raccoon dogs, their burrows and excrements, were found in rare 
oak-hornbeam forests with ground cover of rhododendron, honeysuckle 
and other bushes. They avoid coniferous forests and are only encoun- 
tered there along forest edges, riparian shrubs, or on cliffs along the sea 
coast. In unforested regions, they are met with only in stands of reeds 
along the shore of rivers and lakes. They usually live at an elevation 
of up to 100-300 m above sea level, rarely up to 600-700 m (Bannikov, 
1964). 

In the central zone of the European part of the USSR, they prefer 
deciduous and mixed forests of flood plains, and avoid dense coniferous 
tracts and extensive open bogs and fields. At the northwest, the main 
sites of the raccoon dog consist of southern taiga forest (30% of occur- 
rences in summer and 49.5% in winter), shores of water bodies (26 and 
20.8% respectively), bogs (12.7 and 10.2%), clearcuts and bums (1 1.3 
and 7.1%); in this, mixed forests adjacent to floodplain sections of the 
meadows and fields provide the principal sites both in winter and in 
summer (Geller, 1959). In Novgorod district in flooded forests, the 
number of raccoon dogs is 13; in grassy fir groves, 8; in pine woods, 
1—3 animals per 1000 hectares (Morozov, 1948). In Tatariya in forests. 



94 







11 Fig. 24. Reedgrass and sedge meadows in the lowland of Lake Khanka (southern 
Primor'e). Biotope of raccoon dog. Photograph by A.G. Pankrat'ev. 

raccoon dog encounter frequency is 33% (from 25 in summer to 40— 
60% in winter and spring), in floodplains — ^30% (56% in summer, 

77 10-19% in winter and spring), in ravines with bushes — 11.5%, in 
bogs— 6.2%, in fields— 14.4% (Yu. Popov, 1956). In Voronezh dis- 
trict, they prefer to live in deciduous and mixed forests, especially on 
floodplains where nearly half of all litters were noted; in small glades 
in boggy depressions (more than 1/3 of the litters counted); in large 
tracts of pine, they are confined to forest edges, in bums and clearcuts 
(Obtemperanskii, 1953). In Gor'kovsk district, they prefer the flood- 
plain deciduous forests with rich stands of grass (Kozlov, 1952). In 

78 Byelorussia its favorite habitat is broad-leaved and mixed forest is- 
lands, the copses among meadows and small bogs with dense 
undergrowth of hazel, buckthorn and other bushes, and also dense 
growths of alder thickets and oiser-beds along the banks of rivers and 
lakes; it decidedly avoids dry pine woods (Dol'bik, 1952). 

In Ukraine, it prefers the floodplains with growths of oiser and 
high grass as well as the edges of bogs and flood-pools in the lower 
reaches of large rivers. It establishes itself on river-terrace forests of 
aspen and alder or in oak-groves, especially with dense understory. In 
winter it does not abandon river floodplains (Komeev, 1954). In the 
Donbass, besides floodplains, it willingly colonizes ravines overgrown 



95 



:,v .. . 



-•**''; :jr 




77 Fig. 25. Biotope of raccoon dog — Borzi river valley in its lower reaches, Amursk 
district. Photograph by A.G. Pankrat'ev. 

with shrubs, and in small shrubs in the steppes (Sakhno, 1948). In 
Rostov district, it lives among shrubs, in ravines and depressions as 
well as in growths of reeds (Rail', Kritsksya, 1958). On the shore of 
the Azov Sea, it is found in reed thickets and in growths of shrubs 
(Kostyunchenko, 1950). Similar places are inhabited by this species 
along the shore of the Caspian Sea and in the Volga delta. In the 
northern Caucasus, the raccoon dog lives along the banks of rivers, 
lakes and creeks, in ravines and gullies overgrown with bushes, in 
isolated oak-groves among the steppes and in broad-leaved and mixed 
montane forests. In the belt of montane coniferous forests, they are 
encountered half as often as in mixed and broad-leaved forests 
(Trushchalova, 1959). In Georgia, this species is confined to cut-over 
forests and shrublands along river banks (Meladze, 1947); in Kirghizia, 
they live along swampy shores of lakes, rivulets and in berry-bushes 
and reeds (Berens, Yanushevich, 1956). The raccoon dog was intro- 
duced into Siberia, but quickly died out; it inhabited forested floodplains 
of rivers where nut-pine is found and shrubs are abundant; in summer, 
they frequently occupied open nut-pine woods in searching for food 
(Nadeev, 1940). 

Apparently, mixed and deciduous forests, alternating with small 
open areas near water, represent the most favorable habitat of the 



96 

raccoon dog in its natural range, as well as in places where it has been 
successfully acclimatized. In the majority of northern and central re- 
gions of acclimatization, the raccoon dog prefers, in the snowy period 
of the year, to live in open places (fields, meadows, marshes) where it 
can move easily. 

Food. According to the constituents of its food, the raccoon dog is 
an omnivorous animal. The following moфhological characteristics attest 
to its omnivory: weak development of canines and camassial teeth, flat 
surface of molar teeth, and relatively long intestine (1.5—2 times longer 
than that of other species in the family). The diversity of food objects 
is very great. Animal food appeared in all stomachs and feces, while 
plant food was, on the average, 64%. 

Among animal foods, the most significant frequency of appearance 
(in %) in all seasons and regions on the average are: 



Insects 


47 (maximum, to 93) 


Mouse-like rodents 


46-(85) 


Amphibians 


18-(65) 


Birds 


13-<23) 


Fish 


9-Ч24) 


Reptiles 


8-(36) 


Molluscs 


7-(47) 


Carrion 


M13) 


Insectivores 


5414) 



The absolute significance of one or another food is not only in terms 
of frequency but also volume. In total, the role of mouse-like rodents 
is considerably greater than that of insects. For example, the frequency 
of occurrence of mouse-like rodents and insects in the food of the 
raccoon dog of Voronezh preserve constitutes 60 and 46% respectively. 
The volume of food ("food coefficient" — the product of the number of 
occurrences multiplied by the volume in percentage, divided by 100) 
79 for mouse-like rodents is equal to 58, and for insects — 18 (Ivanova, 
1962). In all regions inhabited by the raccoon dog, it may be confirmed 
that mouse-like rodents occupy the first place in the nutrition of the 
species. The significance of other food groups is very variable and 
depending on the season and the conditions of the regions, insects or 
amphibians, birds or plant food constitute the food base. In one way or 
other, all the above-mentioned food groups are always the main com- 
ponent of raccoon dog food. 



97 




Fig. 26. Change in food of the raccoon dog by seasons (from Yu. Popov, 1956). 



80 



Mouse-like rodents in the food of the raccoon dog are mainly 
represented by voles. Among the latter, those species living in damp 
brushy undergrowth, banks of water bodies and coppices are more 
frequent than other species; these are: reed voles, root voles, water 
voles, common voles and others. In forest regions, the beasts rarely 
feed on common redbacked voles. Mice are 5—10 times rarer than 
voles; as an exception (in Astrakhan district) gerbils are of great im- 
portance in the nutrition of raccoon dog. 

bisects constituting the food of the raccoon dog are more frequently 
represented by: dung beetles, cockchafers or water beetles and water 
scavenger beetles; other groups are more rarely found: tiger and ground 
beetles, carrion beetles, grasshoppers and others. Frogs are more com- 
mon than the other amphibia among the food of raccoon dog. In some 
instances, for example in Voronezh region, fire-bellied toads [Bombina] 
were the usual food (Ivanova, 1962), while in the Ukraine, their usual 
food was the spade-foot toad [Pelobates] (Komeev, 1954). 

Among birds in the stomachs and feces of raccoon dog, the most 
frequent are the remains of either ducks or passeriform birds; com- 
monly passeriforms nesting in the area (buntings, pipits, warblers and 
others) or else migrant species — thrushes, jays, etc. Where there are 
colonies of nesting terns and gulls, their eggs and nestlings are common 
food of raccoon dog. In the northern regions of raccoon dog acclima- 
tization, grouse are usually its prey (Sorokin, 1956). In the Ussuri 
territory, cases of destruction of pheasant nests were recorded. 

The raccoon dog usually utilizes only large fish which are cast up 
in spring after freezing; it rarely catches fish in passage during the 
spawning migration. In Primor'e, it usually collects fish and inverte- 



98 



90 

80 

70 

60 

50 

йО 

30 

20- 

10- 



П 



п 



ш 



12 3^56 
Spring 



тп 

1 2 ЗА 5 6 

Summer 



п 



123^56 

Autumn 



i 23U5 б 

Winter 



79 Fig. 27. Seasonal variation of food groups in the nutrition of raccoon dog (in 
food-units) (from Ivanova, 1962). 
1 — Mouse-like rodents; 2 — Carrion; 3 — Birds; 4 — Amphibia; 5 — Insects; 

6 — Plant food. 




80 Fig. 28. Raccoon dog searching for food on river ice. "Kedrovaya Pad' " preserve, 
southern Primor'e. March 1964. Photograph by A.G. Pankrat'ev. 



brates on the sea shore. It catches small fish from water bodies drying 
out in summer. Snakes are the more frequent prey among reptilians. 
Lizards are 2—3 times rarer. In the southern regions of their acclima- 
tization, the raccoon dog willingly catches young tortoises and digs out 
the clutches of tortoise eggs (Rukovskii, 1950). Both water molluscs — 



99 



7000- 

(д 6000 
i 5000 

О) 

1000 
О 




/^ Adult males 

— ^ Adult females 
~~ Juveniles 



X 



7ШШШЕ1ЛМ I Л Ш Ш 
Months 

80 Fig. 29. Change in weight of the raccoon dog during the year (after Yu. Popov, 

1956). 



bivalves, pond snails and flat-coiled snails — ^and land snails are in- 
cluded in the food of raccoon dogs. 

Among insectivorous animals, the usual prey are shrews and hedge- 
hogs, rarely moles and desmans. In the Far East, the Ussuri mole 
[Mogera robusta] is the usual prey of raccoon dog, a fact which is 
probably determined by the surface position of its feeding burrows. 

Plant food is also highly variable. It may be the vegetative parts 
of plants, bulbs, rhizomes, grains of agricultural crops (most frequent 
are oats, millet, maize), nuts, fruits, berries, grapes, melons, watermel- 
ons, pumpkins, tomatoes, etc. In Ussuri territory, the most important 
are fruits of wild fruit trees, Amur cork trees, acorns and wild grapes. 
Fruits of [cultivated] fruit trees and grapes have a great importance in 
the feeding of the raccoon dogs acclimatized in the southern regions. 
In the northern regions oats, berries (bilberry, cranberry, strawberry, 
and others) and the vegetative parts of plants play a great role. 

The fairly evident seasonal change in the food of the raccoon dog 
is greatly determined by the different availability of these or other food 
items during the year. Thus in late autumn and winter, mouse-like 
rodents appear to be the main food of the raccoon dog, or in the north 
it might be carrion and feces; since amphibians, reptiles, molluscs and 
insects disappear. In the southern regions, the most important food 
items are the grains of agricultural crops, fruits, and also those am- 
phibia not undergoing hibernation (Komeev, 1954; Trushchalova, 1959 
and others). 



100 

In spring, especially during snow melt, the number of mouse-like 
rodents available to the raccoon dog shaфly increase because of the 
poor protection afforded by their burrows. During this period, amphib- 
ians play also their greatest role. During the flight period of cockchafers 
and the reproduction of dung beetles in the late spring, these insects are 
also shown to be main foods. Vegetative parts of plants are very im- 
portant. In summer the food is most diverse and, besides mouse-like 
rodents, the relative role of which decreases, birds are of great impor- 
tance (especially in the nesting period). Insects are important, and fish, 
reptilians and molluscs are found more frequently than in other seasons. 
By the end of summer, a greater role is taken by berries, grains, fruits 
and vegetables. 

In autumn, plant food is the most important item in comparison to 
all other seasons. In connection with the increase in the number of 
mouse-like rodents, their role increases, but the insects, amphibians, 
reptiles and birds gradually disappear from the food ration (Table 1). 
82 The importance of one or another food group differs remarkably 

according to the year. Thus, in the food of raccoon dog, the frequency 
of occurrence of the main food group — mouse-like rodents — may change 
nearly two-fold (Table 2). The decrease in the role of mouse-like ro- 
dents in the food (determined by the drop in their number) entails a 
shaф increase in the significance of birds, amphibians and insectivores. 
In other instances, the lack of mouse-like rodents is compensated by the 
intensive eating of carrion (Novgorodsk and Kalininsk districts; Geller, 
I 1959), feces (Tatariya; Yu. Popov, 1956), or fruits (Caucasus; 
Trushchalova, 1959). Parallel to the decrease in the role of mouse-like 
rodents in the food of the raccoon dog, the general significance of plant 
food also decreases. 

The food contributions of animals inhabiting different places also 
vary somewhat. Thus, mouse-like rodents (common voles) are highly 
important in the food of raccoon dog litters at the forest edges in 
Voronezh preserve. Because of the low number of mouse-like rodents 
within the forest, their importance in the food of litters living in the 
center of the forest was very low; the deficiency of rodents here is 
compensated by amphibians and insects (Table 3). The food of litters 
living in floodplains was more diverse; the most important were water- 
fowl (Ivanova, 1962). Differences nearly as shaф as these in the food 
components of raccoon dogs inhabiting various biotopes were noted in 
Tatariya (Yu. Popov, 1956), in Belorussia (Samusenko and Golodushko, 
1961) and in other regions. 



101 



Table 1. Seasonal change in raccoon dog food in Primor'e 

(E.P. Spangenberg) and in the northwestern European part of the USSR 

(frequency, %; Geller, 1959) 



^\.,^^ Region, season, 
^\^number of data 


Primor'e 
Spring and 


territory 
Autumn, 


Northwestern USSR 
Spring and Autumn, 


Type of food ^^^^^ 


summer, 

97 


26 


summer, 

214 


130 






Animals 






Insectivores 







— 


5.6 


7.5 


Mouse-like rodents 


18.6 




15.4 


17.7 


48.3 


Birds and their eggs 


8.3 




— 


16.8 


4.1 


Lizards 


7.2 




— 


— 


— 


Snakes 


4.2 




— 


Trace 


— 


Turtle eggs 


5.2 




— 


— 


— 


Amphibia 


65.5 




15.4 


52.8 


19.1 


Fish 


18.6 




23.0 


1.3 


10.0 


Insects 


32.0 




23.0 


1.3 


10.0 


Molluscs 


47.3 




11.3 


Trace 


10.0 


Carrion 


2.1 




— 


6.5 


15.8 


Feces 


— 




— 


5.7 


1.6 






Plants 






Nuts and grains 


11.3 




11.3 








Grass 


26.8 




46.1 


57.2 


— 


Oat 


— 




— 


3.7 


68.3 


Rice 


4.2 




— 


— 


— 


Polygonum, millet, com 


2.1 




19.2 


— 


— 


Berries 


2.6 




— 


6.6 


10.7 


Grapes 


— 




46.1 


— 


— 


Apples 


— 




62.2 


— 


— 


Cork tree 


— 




61.5 


— 


— 


Acorns 


— 




— 


— 


5.0 



The sharp differences in the importance of one or another food in 
different seasons in various years and in various habitats make it 
difficult to establish rules governing the variability of food components 
(Tables 1, 2, and 3a). 

In northern and middle latitudes, mouse-like rodents occupy first 

place among animal food; in the south and west — ^insects. As one 

moves southward, the role of birds and molluscs increases. Naturally, 

83 geographic variability of food includes replacement of some food types 

by others within one food group (different species of mouse-like 



102 



Table 2. Change in importance of foods of raccoon dog in various years 
(frequency of occurrence, %) 



\ч Region, 


author, 


Voronezhsk 


Tatariya 


Northwestern 


\. year 


district 


(Yu. Popov, 


European 






Ф 


/anova. 


1956) 


part of USSR 








1962) 




(Morozov 
Geller, 


, 1956; 
1959) 


Type of food 
















^ 1958 


1959 


1949 1950 


1941 


1947 


Insectivores 




5.4 


15.0 


No data 


0.9 


6.9 


Mouse-like rodents 


74.6 


49.7 


59.3 76.0 


40.3 


22.3 


Birds 




13.5 


33.2 


No data 


4^3 


10.2 


Reptiles 




2.2 


3.2 


No data 


— 


— 


Amphibians 




12.4 


23.7 


14.1 6.5 


11.9 


29.8 


Insects 




48.6 


43.9 


56.0 53.0 


29.8 


7.6 


Plants 




50.8 


40.8 


40.3 28.6 


94.7 


34.0 



rodents, insects, berries and so on in different regions). The extremely 
scanty material for the feeding of raccoon dog in the Far East does not 
permit any [comparative] analysis of changes in food components of 
acclimatized animals [elsewhere] [the changes mentioned by Geller 
(1959) are confined to annual and seasonal variability of feeding]. 

Home range. Data about the size of the home range are almost 
absent. The distance between the inhabited burrows, which may serve 
as an indirect indication of the size of the home range, is extremely 
variable. The burrows may be separated from each other by some 
kilometers or (as in Primor'e and on Askol'd Island) very close to each 
other. Burrows were distributed in colonies consisting of Ф-5 (up to 10) 
each in Rostovsk district where the foraging area of the raccoon dog 
reaches 10 km^ (Stepanov, 1939; Rail' and Kritskaya, 1953). 

In the northwestern regions of acclimatization, judging by 19 tracks, 
the length of the daily travel of the raccoon dog is equal to 3—6 km in 
winter, in early spring 1 5—20 km, in late spring 6—8 km, in summer 6- 
10 km, in early autumn 8—12 km, and in late autumn 2—3 km. The area 
within which these travels take place, was equal to 1 to 12 km^ (Geller, 
1959). In Tatariya, the length of the daily travel of the animals in 
search of food in early spring reaches a maximum of 10—14 km; in 
summer, when food is abundant, this travel is less and the radius of 
activity of seven raccoon dogs consisted of 600-800 m. In autumn, 
when fat is accumulating, the length of the foraging circuit may reach 
8 km, but usually is 4—6 km. In winter, during periods of thaw, they 



103 



82 Table 3. Foods of raccoon dog litters in different habitats in Voronezh 
Preserve (frequency of occurrence, %) 



^^---^,_^ 


Biotope 


Floodplain 


Forest 


Forest edge 


Type of food 


^^^^ 


(burrow No. 


1) (burrow No. 


2) (burrow No. 3) 


Mouse-like rodents 


25.7 


5.0 


56.0 


Birds 




48.6 


— 


20.0 


Reptiles 




8.6 


10.0 


8.0 


Amphibians 




31.4 


70.0 


16.0 


Insects 




62.8 


100.0 


52.0 


Plants 




25.7 


20.0 


56.0 



do not go further than 100-150 m away from shelter (Yu. Popov, 
1956). 

Burrows and shelters. Raccoon dogs use several types of shelters: 
beds and lairs in open areas, and burrows, both temporary and perma- 
nent ones (breeding and wintering dens). 

In the Ussuri territory, summer beds are generally places on dry 
slopes under a protective rock, boulders or in brushy undergrowth. In 
the Khankaisk lowland, beds/nests found in September— October were 
between hummocks on a thick (4.5-6 cm) layer of accumulated cuttings 
of dry sedge, reed and other grassy vegetation. At the bottom of the 
nest, scraps of wool were found. The internal diameter of such nests is 
84 78—100 cm, the depth is 32—36 cm. Around the nest the reeds usually 
form an obstruction by arching above the nest. In the Ussuri territory, 
beds are found in the depressions between roots of old tress and in the 
spacious hollows of fallen lime trees [Tilia]. 

Based on investigation of 49 burrows in the Ussuri territory, the 
majority of the holes are located 500-800 m away from the banks of 
water bodies, most frequently (41%) at eastern points of the compass. 
Slopes on which burrows were found were covered with forests (88%) 
or shrubs (12%). 78% of the burrows were found under piles of stones, 
12% in loose soil, 8% in rock niches, and 2% under tree-roots (Bannikov 
and Sergeev, 1939). 

The horizontal diameter of the passage of burrows beneath stones 
is often equal to 24—29 cm (2 1 to 3 1 cm), vertical diameter 20—24 cm 
(20 to 34 cm). In the burrow there are usually 1—2 entrance holes, in 
some instances 3—5. The longest passage to the nesting cavity is 2 m, 
more frequently 1.5 m, i.e., the burrows of this type are comparatively 
short and simply constructed. The nesting cavity has a flat bottom and 
a dome-shaped roof; its dimensions are 50-70 x 30-45 cm. 



104 



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105 




84 Fig. 30. Burrows of raccoon dog. Scheme of location and cross section (after 

Bannikov and Sergeev, 1939). 

Most often (36 burrows out of 38) there is only one entrance to the 
cavity; the side burrows join together in front of the nesting cavity in 
one principal passage. The bottom of the cavity is usually lined with 
different dry plant remains, leaves of hazel, oak, maple, lespedeza, 
stems and leaves of grasses and herbs. 

Near the burrow, usually not farther than 10-15 m, and in the 
burrows with litters, toilets in the form of pits are located, sometimes 
directly near the entrance, filled with excrements. Near the burrow 
under piles of stones occur latrines in the form of numerous heaps of 
excrements on flat rocks. In such cases, the latrine is frequently located 
20-30 m, or farther, from the burrow. 

Burrows excavated in soft ground in Ussuri territory were situated 
along the slopes of river valleys, in places poor in rock outcrops. They 

85 are quite simple, having 1-3 exits averaging 21 x 24 cm in diameter 
(18-22 X 25-29 cm). Fan-shaped scattered soil and gravel are found 
in front of the entrance to the hole and lower along the slope. The 
burrow passage usually passes downward directly into the ground without 
any turns at an angle of 20-25°; its length reaches 6 m (most frequently 
3 m) and ends with a nesting chamber which does not essentially differ 
from that described above in burrows under stones, but in the holes dug 
in soft ground, the layer of leaf and grass bedding is less. In some 
burrows a blind side hole proceeds from the nesting chamber which 
sometimes rises slightly upwards, in which the concealed animal is 
found when the burrow is dug up. Burrows in rock niches are only 
accessible for external inspection. There is one entrance to such bur- 
rows; wide fissures (more than 50 cm) and those opening upwards are 
not occupied by raccoon dogs (a burrow located under the roots of an 
old lime tree had 3 entrance openings under the tree). 



106 



Entrance No. 4 



Chamber* 
of the burrow 





Bifurcation No. 1 
Light-gray loam 



Entrance 
No. 1 



Loam with / / sandi 



84 



3.5 и 



3.3 



If. 6 



Fig. 3 1 . Scheme of construction of raccoon dog burrow (from Yu. Popov, 1 956). 
*ln Russian original hotel, which means "caldron, boiler, pot" — Sci. Ed. 



Shelters of the raccoon dog in places of their acclimatization are 
similar to those described. In Tatariya and Rostovsk district, summer 
beds, in the form of small holes without bedding are situated on dry 
places at the base of slanting shrubs growing in depressions, and the 
shores of marshes and lakes (Rail' and Kritskaya, 1953; Yu. Popov, 
1956). The raccoon dog sometimes constructs a lair in tall, dense sedge 
thickets in the form of a plaza (70 x 100 cm) on small piles of mowed 
sedge or other grass remaining from the previous year. In Tatariya, 
winter lairs on the frozen swamps in dense growths of reed and cattail 
(Yu. Popov, 1956) resemble lairs in the Khankaisk lowland. Raccoon 
dogs accumulate the dry sedge to serve as bedding, while the surround- 
ing high grasses bend under the weight of the snow forming an arch 
above the lair. Below it the animals construct a whole labyrinth of 
hidden passages, ending in an outlet on the surface of the snow cover, 
sometimes 15—20 cm away from the lair. Such lairs constituted in 
Tatariya riiore than 37% of all aboveground shelters. 

In many regions of acclimatization, typical lairs are under hay- 
stacks and shocks. Such dens are usually also used in winter or for 
whelping; they are situated under tangled fallen trees, under stones, in 
old sheds, in trenches, in dug-outs, and in entrenchments remaining 
after the war. In such dens, there is bedding of dry grasses and leaves. 
Shelters in the hollows of fallen trees or in hollows located at ground 
level are quite frequent. Thus, in the northwestern European part of 
USSR, 17% of winter dens of raccoon dog were found under the roots 
of trees, 13% — ^in dug-outs, 10% — in entrenchments, 3% — in hollows 
and stacks (Geller, 1959). In Tatariya, 5.9% of the lairs were found in 
hollows (burrows not included) — 4.8% under haystacks and 5.3% — ^in 
heaps of dry branches (Yu. Popov, 1956). 



107 

The significance of burrows as shelters for the raccoon dog in 
places of acclimatization may be highly variable even in neighboring 
regions. Thus, in Voronezhsk district, about 75% of the litters of the 
raccoon dog were found in burrows (Obtemperanskii, 1952), in Voronezh 
Preserve — more than 86% (Ivanova, 1962). In the latter, old burrows 
of badgers and red foxes are especially numerous, and the ground is 
sandy. In the northwestern regions, about 50% of the raccoon dogs 
occupy burrows (Geller, 1959); in the Ukraine, they rarely utilize 
burrows (Komeev, 1954), but in Rostovsk district they live, as a rule, 
in burrows (Rair and Kritskaya, 1953). 

The raccoon dog usually hurries to occupy the old burrows of 
badgers and red foxes, and digs itself a new burrow only when there 
are no free ones. In Tatariya, 77.8% of the burrows occupied by the 
raccoon dog were old ones of badgers (38.6%) or of red foxes (39.2%) 
(Yu. Popov, 1956). The same phenomenon was observed in other re- 
gions. In using the burrows of badgers and foxes, the raccoon dog, as 
a rule, does not dig additional side burrows and does not construct a 
new nesting chamber, but only brings in dry bedding, throwing away 
the rotted. On occupying the complicated holes of badgers, the raccoon 
dog uses 2—3 side burrows and the others are neglected. 
86 Many cases of wintering of raccoon dog and badger in one hole 
have been recorded. This is possible, because the badger enters into 
hibernation at least 2 weeks earlier than the raccoon dog and leaves the 
burrows 2 weeks after the raccoon dog. In the event that the raccoon 
dog remains longer in the burrow, the badger drives it away and not 
rarely bites it to death (Komeev, 1954; Geller, 1959; Ivanova, 1962 
and others). Exceptionally, cases of cohabitation of litters of badger 
and raccoon dog in one burrow are known (Tikhvinskii, 1938; Yu. 
Popov, 1956 and others). 

In those cases, when the raccoon dog itself digs, it constructs 
temporary as well as permanent burrows. Temporary burrows are simple 
in construction, small and short (length to I m), always with a single 
entrance hole. The animals dig them in soft ground, often under bushes 
or young pine trees, usually on precipitous slopes, along river banks, 
at forest edges and in fields. Permanent littering and wintering burrows 
are more frequently dug on the slopes of forest ravines, in floodplains 
of rivers and at the borders of bogs, not far from forest edges or 
openings in the forest and often on elevated parts of watersheds. In this 
case, preference is shown for deciduous and mixed forests of various 



108 

species, with a rich forest understory. The nature of the ground is not, 
apparently, of decisive importance, but sandy soils are preferred. 

Proper burrows in Voronezh Preserve had one entrance hole in 
57.1% of cases, two in 25% and more than 2 entrance holes in only 
18% (Ivanova, 1959). This relationship of burrows with various num- 
bers of passages is also found in other regions. The general length of 
the burrow ranges from 1.5 to 5 m, often 2.5—3 m. A chamber 40-50 
cm in diameter is situated at a depth of 60-90 cm (usually 70-75 cm). 
Width of the entrance hole exceeds its height, its dimensions being 
24x29 cm (from 19 to 41 cm). One passage usually leads to the 
chamber; as an exception, blind side holes occur, proceeding 60-80 cm 
away from the chamber. The floor of the chamber is lined with a small 
layer of bedding made of dry grass and leaves. 

At the burrow, soil thrown out in a fan shape over 1.5—1.8 m 
occurs. The dug-out soil and the area around the burrow are trampled 
down by the animals, and evident traces of their thin claws are usually 
seen on the walls of the burrow. 5—10 m, but sometimes 2—3 m, from 
the burrow is located a latrine with pits. The trails of the animals are 
seen on the discarded soil but these paths are narrow (not exceeding 20 
cm), and this distinguishes the holes of raccoon dog from those of 
foxes, the paths of which are broad (25-35 cm) at the burrow, and the 
excavated soil coming from the hole is rounded in shape. The burrow 
of the raccoon dog differs from those of the badger in that the soil 
thrown from the hole of the latter is elongated, and the trails are not 
usually seen, since all the excavated soil is trampled down by the feet 
of the animals (Yu. Popov, 1956; Ivanova, 1959, 1962 and others). 

The raccoon dog, apparently, changes the shelters seasonally. Thus, 
in Tatariya, the beasts overwinter in burrows, dens or winter beds. 
With increased activity in March, they often start to use temporary 
burrows or beds, but at the end of April, before whelping, they again 
occupy permanent holes, throwing out the old bedding from them and 
bringing in new. By the middle of June, when the pups are about one 
month old, the animals leave the littering shelters and use temporary 
summer shelters, only rarely visiting the burrows. In autumn, visits to 
the permanent holes become frequent, and in the middle of October, the 
animals clean them a second time and change the bedding; at the end 
of October, they finally settle in their winter shelters (Yu. Popov, 
1956). 

Daily activity and behavior. The raccoon dog is a nocturnal and 
crepuscular animal. However, in the short nights of summer, activity 



109 

occurs during sunny periods in the morning and evening hours. During 
the day, activity of the animals is curtailed and they hide in shelters or 
spend the time bedded down. During the period of rut, pregnancy and 
the initiation of nursing the young, from March to May, the raccoon 
dogs are very cautious and are active almost exclusively in the dark 
period of the 24 hours. In summer, when the pups begin to feed inde- 
87 pendently, they frequently are encountered during the daytime. In autumn, 
in September, regardless of increased feeding, raccoon dogs rarely 
come out in the daytime and usually leave their shelters only at twi- 
light. In winter, the animals are active at twilight and during the night. 

Upon leaving the burrow the raccoon dog in search of food follows 
the shore of the nearest water body. It frequently goes into water and 
wanders along the shallows. 

On the sea coast, along shores of lakes and rivers, at dawn and in 
the evening, it is almost always possible to see feeding animals. In the 
forest the raccoon dog moves slowly, looking under every bush, pile 
and stump. Only when crossing openings does it move at a quick trot. 
The raccoon dog is cautious but in case of danger it does not run away, 
but conceals itself, camouflaging itself very skillfully. If they are over- 
taken, some animals (in the Far East) pretend to be dead, but on the 
first opportunity, they run away and conceal themselves nearby. The 
entire litter usually grows fat together at the end of summer. It is 
possible to observe two or three adult animals, feeding in immediate 
proximity to one another. 

Having comparatively short legs, the raccoon dog, in the presence 
of a snow cover 10-12 cm deep, already stops "dragging", and searches 
for food with difficulty. In a snow depth of more than 20 cm deep or 
deeper the animals move with great difficulty, "swimming" in the snow. 
The weight load for the track is relatively great: in December it is 75— 
80 g/cm^, in March 33—37 g/cm^ (Yu. Popov, 1956). The sense of smell 
is very sharp, and hearing and vision are more weakly developed than 
in closely-related species. 

Hibernation, winter sleep. True winter toфor, i.e. deep uninter- 
rupted sleep, associated with a 8Ьаф fall in metabolic level and body 
temperature, is not observed in the raccoon dog. However, the general 
metabolism may decrease by about 25% (Sokolov, 1949). 

In warm winters in the Ussuri territory, as well as in places of 
acclimatization (Ukraine; Komeev, 1954), the beasts do not go into 
hibernation during the course of the entire winter and only during 
snowstorms they do not go out of their shelters for some days. Usually 



no 

in December, when the snow cover reaches 15-20 cm, the activity of 
the raccoon dogs shaфly decreases and they leave their shelter for a 
short time only on warm, quiet days and they do not go farther from 
the burrow than 150-200 m. From February to the beginning of March, 
the activity of the animal again shaфly increases, both from food 
deficiency as well as the beginning of rut, and this period leads to an 
increase in the movements of the raccoon dog. During spring snowfalls 
(March— April) the animals again may go into shelter for some days. 

At the beginning of the winter period, the raccoon dog puts on an 
average of 18—23% of subcutaneous and 3—5% of internal fat (of total 
weight). Animals putting on less than this amount (though often gain- 
ing) as a rule remain wandering and rarely survive the winter. During 
the first years of acclimatization, when animals were not yet suffi- 
ciently adapted to catching the local food, as a result of this they 
exhibited weak fattening in autumn, and frequently did not hibernate 
(Obtemperanskii, 1958 and others). Low temperature exerts little influ- 
ence on the activity of the raccoon dog. Instances are known of animals 
being encountered at -20 to -25°C. 

Raccoon dogs kept in artificial burrows in hutches fell into hiber- 
nation in December, but aroused and fed during a time of warming; in 
the period of rut (in mid-February) they wandered about, but then at 
several times hibernated again (Obtemperanskii, 1958). Confirmation 
of partial winter activity of the raccoon dog is attested by their being 
caught by hunters. Thus, in Tatariya, about 80% of the animals are 
caught from November to January and 20% from the 1 st of January to 
the 15th of February (Yu. Popov, 1956). In the northwest of the coun- 
try, about 37% of the animals are caught in December, in January — 28% 
and in February— 18% (Geller, 1959). 

Seasonal migrations and transgressions. The raccoon dog is a 
sedentary animal, and regular seasonal migrations are not known for 
this species. 

In the Ussuri territory, hunters have reported migrations of raccoon 
dogs in unfavorable years. In the Lake Khanka depression, massive 
migration of the animals to China seemed to have been observed in the 
autumn of 1929 when, within a few days, up to one hundred animals 
were seen moving in one direction. Such migrations are doubtful; prob- 
ably what occurs is the migration in the winter of a small number of 
animals living during the summer in reeds from flooded depressions to 
dry places. 



Ill 






tii3L^.l jJ^ -- 



Fig. 32. A young raccoon dog, just appearing after winter sleep. "Kedrovaya Pad" 
preserve, southern Primor'e. 17 February 1965. Photograph by A.G. Pankrat'ev. 

After releases in new regions, there were migrations of individual 
animals for 20-80 km from the place where they were originally intro- 
duced (Rair and Kritskaya, 1953; Trushchalova, 1959 and others). 
There is the possibility of more distant migrations, since intrusions of 
the animals onto the tundra have been recorded (Skrobov, 1958). An 
emigration of about 600 km was recorded in Ukraine (Komeev, 1954). 
The greatest migrations of the raccoon dog are recorded in floodplains; 
during the final spring floodwaters, females with litters move up to 
divides. 

Reproduction. Rut is observed from the beginning of February to 
the end of April depending upon the region and climatic conditions. 
Later spring may delay the beginning of rut for 2—3 weeks. The return 
of cold weather, and especially snowfalls, usually interrupt rut. In the 
Ussuri territory, rut takes place on the average in the middle of March, 
but it often is delayed until the end of March or beginning of April. In 
the Ukraine, in Voronezhsk district and Tatariya, rut occurs in the 
middle of February (Komeev, 1954; Yu. Popov, 1956; Obtemperanskii, 



112 

1958); in the northwest, in the beginning or middle of March (Geller, 
1959 and others). Rut continues for 2-3 weeks, but in captivity, it is 
usually not less than 3 weeks, and often 26-28 days, and during food 
shortage, up to 40 days (Pavlinskii, 1937). 

Raccoon dogs are monogamous animals and pairs are formed even 
in autumn, usually in October or November. Fights among males occur 
occasionally at the time of rut, are brief and not violent, and are 
89 accompanied by yelping and hollow growling. In captivity, polygamy 
is possible when one male is in contact with 4—5 females. Copulation 
occurs most frequently during the night or early in the morning, usually 
in quiet weather. Coitus lasts for 6-9 minutes on the average (from 2 
to 26 minutes). Estrus in females continues from several hours to 6 
days and during this period they mate up to 5 times, usually 2-3. After 
20-24 days, estrus is repeated even in pregnant females (Petryaev and 
Khatkevitch, 1931; Pavlinskii, 1937). 

Pregnancy lasts 61-70 days (often 59 days [sic]*). Young are bom 
in April, but mostly in May. Cases are known when the main mass of 
births is delayed until June, and in a few cases newborn have been 
found as late as September. 

In the Far East, the litter mostly consists of 6-7 young, but their 
number may reach 15-16 (Firsov, 1929; Przheval'skii, 1870**). The 
size of the litter in the regions of acclimatization fluctuates greatly, in 
connection, probably, with fluctuation of conditions in various years. 
Thus, in Voronezhsk district (Obtemperanskii, 1958) average litter size 
was 4.9 pups (from 4.3 to 5.75 in different years); in the northwestern 
districts in 1938-1939—5.2; in 1947—5.9; in 1946-1949 average, 6.3 
(Morozov, 1953; Geller, 1959). In Tatariya (Yu. Popov, 1956) the 
average litter size was 7.3 pups; in the Ukraine (Komeev, 1954) — 7.9 
(to 16 pups); in Lithuania — ^from 4 to 13, average, 9.5 (Prusaite, 1961). 
The average occurring in captivity is 7.1 pups (Petryaev and Khatkevitch, 
1931). Females giving birth to pups for the first time have smaller 
litters than females of older age, and their litters are 1 or 2 pups fewer. 
The ratio of sexes at birth is nearly 1 : 1 ; but males are somewhat more 
frequent. Thus, in Tatariya (Yu. Popov, 1956) litters contained 50.9% 
males and 49.1% females; in the northwest (Geller, 1959) — 51.7% and 
48.3% respectively; and in Voronezhsk district (Obtemperanskii, 
1958)— 58.7% and 41.3%. In State animal farms (Povlinskii, 1939 
52% males and 48% females. 

*Russian original inconsistent — Sci. Ed. 

**In Russian original, 1947 (not in Lit. Cit.) — Sci. Ed. 



113 

Barrenness of females in different regions of acclimatization con- 
stitutes about 12% (Yu. Popov, 1956; Geller, 1959 and others). In 
captivity, after one mounting [by male] 30.9% of the females remain 
non-pregnant and after three mountings, practically no females are 
barren (Starkov, 1940). The male takes an active part in raising the 
young. In Tatariya the average number of pups in the litter decreases 
from 7.3 in May to 5.9 in June due to mortality (Yu. Popov, 1956). 
Juvenile mortality increases in June when the pups leave the shelter (at 
an age of 25—30 days), and reaches its maximum in July and August 
when the growing pups separate from the litter. The litter finally breaks 
up at the end of August— September, and in October juveniles, reaching 
the size of adults, unite in pairs. During this period about 35% of the 
young have died, and each pair of adults during this time produces 4 
juveniles (Morozov, 1953). By September and October, about 50% of 
the young have died (Geller, 1959). 

Growth, development, and molt. Young are bom blind, covered 
with short, dense, soft wool, without guard hairs, of dark-slaty, nearly 
black color. Weight of newborn, 60—1 10 g. Males weigh 5-10% more 
than females. Eyes open on the 9th— 10th day. On the 14th— 16th days 
teeth erupt — beginning with the upper canines, then the incisors and 
lower canines. Approximately from the age of 10 days, guard hairs 
begin to grow on the hips and shoulders, and then near the ears and on 
the cheeks. The dark color of the pup gradually becomes gray, starting 
with the abdomen, 2 weeks after birth. Guard hairs start to grow 
especially quickly at the age of 1 .5—2 months when the black or black- 
ish-brown color prevailing in the color of pups quickly changes to gray. 
The black tones remain only around the eyes. On all the interorbital 
area, fi-om the forehead to the nose, the brown color of the underfur 
also changes into light gray. By this time, as a result of lengthening of 
hairs on the cheeks, "side- whiskers" begin to form. In 3 -month-old 
90 pups, the guard hairs are shorter than those of adults only on the chest, 
chin, tail and distal parts of the extremities. 

Lactation lasts for 45—60 days, but from 3 weeks or one month of 
age, the pups start eating frogs, insects and other food brought to them 
by the parents. At the age of 1 month, the pups weigh 550-650 g; at 
2 months, 1 100-1300 g; 3 months, 2500-2900 g; and at 4 months — 
about 4 kg. At an age of 4.5— 5 months, young attain the weight and size 
of adults and are almost not different from them in color (Pavlinskii, 
1937; Komeev, 1954; Yu. Popov, 1956; Geller, 1959 and others). 

The lowest weight of adults is attained in March (about 3 kg). By 



114 

August— beginning of September, males reach their greatest weight— 
6.5-7 kg (certain individuals, 9-10 kg). Females deposit fat reserves 
more slowly and maximum weight is usually attained one or one and 
a half months later than males, i.e. by September-October. Juveniles 
attain greatest weight another month later than females — in October- 
November. In autumn, fat constitutes 30-35% of live weight of the 
animals and forms a subcutaneous layer up to 1.5 cm (Yu. Popov, 
1956; Geller, 1959). 

Sexual maturity is attained at an age of 8-10 months. The duration 
of life is not known. In nature, individuals older than 6-7 years are 
encountered. In captivity, animals are known to have lived to 1 1 years. 

The age composition of populations is insufficiently studied. In the 
northwestern regions, juveniles constituted 53.8% of the population in 
the hunting season and animals of older ages — 46.2% (Geller, 1959). 
Among animals caught in Tatariya in the hunting season, juveniles 
constitute 57.2% of the population, adults — 42.8% (among them, 19% 
are older than 1 year, 1 1 .3% older than 2 years, 6.7% older than 3 
years, 3.9% older than 4 years, 1.9% older than 5 years). In Lithuania 
in 1957—1959, one-year-old animals constituted 67.5% of the. popula- 
tion (Prusaite, 1961). 

Molt occurs only once a year — ^in spring. Shedding of the underfur 
starts in February— March depending upon the region, coming of spring, 
and condition of the animals. Shedding of underfur starts on the neck 
and withers; then molt extends to the shoulders, back, sides and the 
posterior part of the body. By the middle of May— June (and July in the 
northern regions), guard hair almost alone remains on the skin, and the 
fiir acquires a dark color because of the proximity of the tips of the 
guard hairs, which have black tips. From June onward, guard hairs are 
gradually replaced and the underwool starts growing. Renewal of the 
latter takes place slowly compared with guard hair, and therefore the 
pelage attains normal development from the last days of October, to 
November, and even December. The underfur (new) per 1 cm^ of the 
rump is in August — ^2072, in October — 6264, and in December — ^9624 
(Tmshchalova, 1959). In old males, moh takes place more quickly than 
in females and juveniles (Komeev, 1954; Morozova, 1955; Geller, 
1959 and others). 

Enemies, diseases, parasites, mortality, competitors and popula- 
tion dynamics. Among the enemies of the raccoon dog, the most important 
is the wolf It destroys raccoon dogs early in spring — in February- 
March, and also in summer. However attacks on raccoon dogs by 



115 

wolves have been noted in late autumn also. Stray dogs destroy mainly 
the growing pups. Apparently, during the period of litter dissolution, 
red foxes also destroy pups. Instances are known of foxes biting rac- 
coon dogs to death (see below). 

In Tatariya, out of the 54 recorded cases of death of raccoon dogs 
by predators, 55.6% of the animals were killed by wolves, 27.8% — by 
stray dogs, 11.1% — by red foxes, 3.7% — by the golden eagle and 
1.8% — by the eagle-owl [Bubo bubo] (Yu. Popov, 1956). In the north- 
western regions, out of the 186 recorded cases of death of raccoon dogs 
by predators, 64% were attributed to the wolf, 14.3% — to fox, 12.8% — 
to stray dogs, 6.3% — ^to lynx and 2.6% — to birds of prey (Geller, 
91 1959). Lynx, because of its low numbers, causes little damage to the 
raccoon dog, as observed in the majority of regions; still more rarely 
do brown bears attack raccoon dogs. Among birds of prey involved in 
attacks, usually on pups, are golden eagle, white-tailed eagle [Haliaetus 
albicilla], goshawk and eagle-owl. 

Among the raccoon dogs' competitors, fox and badger are the most 
important, since foxes play a role with raccoon dogs in feeding on 
mouse-like rodents, whereas the badger generally shares with them 
feeding on insects and plants. However in the summer period, thanks 
to the abundance of food, the various habitats and different methods of 
hunting, there is no shaф competition among these species of predators. 
This competition becomes somewhat shaфer in early spring (February, 
March), in the most difficult period of the year. Competition with foxes 
and badgers for burrows is practically absent, for the raccoon dog is 
entirely unassuming in its struggle for shelter (see above). However, 
individual cases have been recorded, when badger or fox killed the 
raccoon dog which had occupied its burrow. 

The parasitic fauna of raccoon dogs was investigated in the Far 
East (Shpringol'ts-Shmidt, 1935; Petrov, 1941), in State animal farms 
(Gusev, 1951; Savinov, 1953 and others), in Tatariya (Yu. Popov, 
1956), in the Ukraine (Komeev, 1954), in the northwestern regions 
(Antipin, 1946; Morozov, 1953; Savinov, 1953; Geller, 1959), in the 
northern Caucasus (Trushchalova, 1959), in the lower Volga region 
(Sviridov, 1952), and in Lithuania (Prusaite, 1961). 

In ail, 32 species of parasitic worms are known from the raccoon 
dog: 8 species of trematodes {Opisthorchis felineus, Clonorchis sinensis, 
Pseudamphistomum truncatum, Euparyphium melis, Metagonimus 
yokogawai, Plagiorchis massino, Alaria alata, Paragonimus 
westermani), 7 species of cestodes {Diphyllobothrium mansoni. 



116 

Taenia hydatigena, Т. polyacantha, Multiceps multiceps, M. serialis, 
Diphylidium caninum, Mesocestoides lineatus) and 17 species of nema- 
todes {Ascaris columnaris, Toxascaris leonina, TfoxocaraJ canis, 
Capillaria yamaguti, С plica, С putori, Thominx aerophilus, 
Ancyclostoma caninum, Uncinaria stenocephala, Molineus patans, 
Crenosoma vulpis, Dioctophyma renale, Physaloptera sibirica, Thelazia 
callipaeda, Strongyloides erschowi, Trichinella spiralis, 
Macracanthorhynchus catulinum *). 

In the Far East, 18 species of parasitic worms were found in 
raccoon dogs, and 12 species were found in regions of acclimatization. 
Such species as Clonorchis sinensis, Metagonimus yokogawai, 
Paragonimus westermani were not found in acclimatized animals due 
to the absence of their intermediate hosts in Europe — the Eastern mol- 
luscs. Some species: Opisthorchis felineus, Plagiorchis massino. Taenia 
hydatigena, T. polyacantha, Multiceps multiceps, M. serialis, Ascaris 
columnaris, Ancyclostoma caninum were acquired by raccoon dogs 
during the process of acclimatization. 

In the northwestern regions there has been noted a high degree of 
infection (up to 96% and 1 0-20 parasites per 1 cm^ of small intestine) 
with the trematode Alaria alata; somewhat less (33.7%) — ^with the 
nematode Uncinaria stenocephala (Savinov, 1953; Geller, 1959). In 
Tatariya, a great number of animals were found to be infected by 
Alaria alata, but the total number of infected animals was less (76.2%) 
than in the northwest (Yu. Popov, 1956), in Lithuania, a lesser number 
of animals (54%) was infected by this parasite (Prusaite, 1961). In the 
Ukraine, 3 species of parasitic worms were found, including Alaria 
alata in 33.3% of animals. Infection with Alaria takes place through 
frogs infected with the metacercaria** of Alaria. In the Caucasus 
(Trushchalova, 1959) a considerable number (27.9%) of raccoon dogs 
are infected with the ascarid Ascaris columnaris. 9.3% of raccoon dogs 
of Lithuania (Prusaite, 1961) are infected with the larvae oi Trichinella 
spiralis. However, the pathogenic significance of parasitic worms to 
raccoon dogs is little studied. In Kievsk district in 1952, there was 
massive mortality among raccoon dogs due to bronchopneumonia of a 
trematode nature (Komeev, 1954). 
92 Six species of fleas have been found on raccoon dogs: Chaetopsylla 

trichosa, C. globiceps, Paraceras melis, Ctenocephalides canis, 

*an acanthocephalan; misspellings in Russian original have been corrected — 
Sci. Ed. 

**Actually, mesocercaria — Sci. Ed. 



117 

С. felis, Pulex irritans, as well as chewing lice, Mallophaga; 5 species 
of ticks: Dermacentor pictus, Ixodes ricinus, I. persulcatus, I. 
crenulatus and the causative agent of mange — Acarus siro*. The latter 
is fairly widespread in the Ukraine, Voronezhsk district, Lithuania and 
a series of other regions, causing massive outbreaks of mange. Raccoon 
dogs are strongly infected with the biting louse Trichodectes canis; in 
the State animal farms of the Far East, this parasite afflicts up to 90% 
of animals (Shpringol'ts-Shmidt, 1935) causing a marked exhaustion of 
the animals. 

Diseases of the raccoon dog are poorly studied. Cases of rabies 
were recorded in the lower Volga (Isakov, 1949), Voronezhsk district 
(Obtemperanskii, 1958), Lithuania (Prusaite, 1961) and in other 
regions. In the Ukraine (Sakhno, 1948; Komeev, 1954) and in Tatariya 
(Tikhvinskii, 1938) cases of massive epizootics of piroplasmosis were 
recorded. In the northern Caucasus, epizootic carnivore distemper was 
noted among raccoon dogs (Trushchalova, 1959); a known case of a 
raccoon dog ill with leptospirosis (N.N. Rukovskii); in State animal 
farms, carnivore plague, paratyphoid, anthrax, and tuberculosis sick- 
ness (Burov, Buzinov, Bannoviskii, Lyubashchenko and others, 1939). 

Epizootics may be an important factor in reducing the number of 
raccoon dogs. "An almost complete destruction of the whole stock" was 
observed as a resuh of an epizootic of piroplasmosis in Kievsk and 
Chemigovsk districts (Komeev, 1954) and in Tatariya (Tikhvinskii, 
1937). A great number of raccoon dogs are lost to mange, which, 
however, does not lead to a massive die-off of the animals, as is known 
for foxes (Prusaite, 1961 and others). The number of raccoon dogs may 
be shaфly reduced as a result of mortality from rabies (Isakov, 1949). 

Among the abiotic factors which cause a shaф decrease in the 
number of raccoon dogs, the spring flooding which coincides with the 
period of reproduction is important. At that time, young die in the dens; 
and, apparently, a significant number of pregnant females are killed 
(Komeev, 1954 and others). In the northem regions, a cause of reduc- 
tion in numbers may be spring snowfall, involving lack of food, death 
due to predators, increase in the number of non-pregnant females and 
lowering of fertility (Yu. Popov, 1956; Obtemperanskii, 1958; Geller, 
1959 and others). There are no concrete data about the magnitude of 
mortality due to different causes. In the flooded areas of the Dnepr, 
Volga and other large rivers, in some years the greater part of popu- 
lation is killed. However, in favorable years, due to the high fertility 

*Mange is also caused by Sarcop tes— Sci. Ed. 



118 

(see above) the population doubles by autumn, and the number of 
raccoon dog is quickly restored. 

The number of raccoon dog is related to food availability. Thus, in 

Voronezhsk district, 3-4 fold fluctuations in the number of this cami- 

93 vore were observed. The fall and rise in the numbers is related to 

changes in the numbers of the principal food — mouse-like rodents — 

in the preceding year and has a 3-4-year cycle (Ivanova, 1962). 

Field characteristics. A stocky animal of medium size, on short 
legs, with small head and pointed muzzle; ears almost hidden in thick 
fur and typical "side-whiskers" on the cheeks. 

When in danger, conceals itself, closely presses itself to the ground 
and, owing to its brown color, merges with the surrounding soil back- 
ground or forest bedding. In case of direct close approach of a human, 
it usually closes its eyes and lies completely still, even when touched. 
During twilight and night, it is more courageous and sometimes tries to 
defend itself It swims well, willingly enters water and can swim across 
wide rivers and lakes. 

In the places inhabited by raccoon dogs, holes dug in search of 
insects may be found. These holes are 5—15 cm in depth and are 




«•.*' 



♦ 



♦t?» 



Fig. 33. Prints of front and hind paws of raccoon dog in silt. On the right, scheme 

of the usual position of tracks during slow movement. Volga delta, 24 August 

1943. Sketch by A.N. Formozov, about 2/3 natural size. 



119 






•'i. 



•!V 



ьЧ» 



»*> 



»'.' 
••?» 



ilt^ 



Ю-/2ст 
.Mi 






•*!b 



7-Wcm 



»!•* 



"-^cm 






30 ~J5 cm 



Quiet gait while 

searching for 

food 



/5 -20 cm 

Quiet gait 

while 
traveling 



r 

25 -JO cm 



Trotting gait 



30-35 cm 



^IV 



j45l 



Leaping gait 



Fig. 34. Tracks of raccoon dog during various gaits (trom Yu. Popov, 1956). 

reminiscent of badgers, but usually wider. A fan of excavated soil (see 
above) is typical of the burrow. 

Tracks are rounded in form, nearly 5 x 5 cm for the front, and 
4.5 X 5 cm for the hind feet. As a rule, the prints of their claws are 
generally easily seen. Unlike foxes, the series of tracks of the ruiming 
raccoon dog is not straight, but zig-zag. A raccoon dog when fattening 
walks slowly, and looks in each pool, hole and under bushes and 
stumps; moving in this way, the tracks of the hind paws do not reach 
the tracks of the front. Having short legs they leave "drags" [marks] 
and "eyes" [holes] on the snow when it is 10 cm deep. In the snow 
cover depth of 25— 30 cm, the animal leaves a deep furrow in the loose 
snow. 

Typical "latrines" in the form of heaps of excrement are found in 

definite places, usually 10-15 cm from the den or on trails. Unlike 

badgers, the raccoon dog does not bury or hide its excrement. Feces are 

94 40-60 mm in length and 20-30 diameter and are composed of 2—3 

pieces. 



120 

Usually silent; only in the rutting period, the males yelp and give 
a hollow growl. (A.B.). 

Practical Significance 

The raccoon dog provides thick, long, strong, but coarse fur. When 
cage reared, the animals also provided about 100 g of wool, of a quality 
somewhat inferior to that of goats. In a series of regions in the Far East, 
the meat of raccoon dog is used by the local inhabitants as food. 
Analysis of the meat (18% protein, 3% fat; more than 4186.8 
joules/kg'") reveals its high nutritious qualities (Komeev, 1954). 

Data about the size of the catch in the past are almost lacking. In 
the eighties of the past century, the trade of raccoon dog in the Primor'e 
and Ussuri territories was "quite developed" (Silant'ev, 1898) and 
occupied an evident place in the fur trade (Nadarov, 1888). The world 
harvest in its entire range, including Japan, North and South Korea and 
China in 1907—1910 comprised 260-300 thousand pelts (Kaplin and 
others, 1955); of which, judging by indirect data, about 5—8%, i.e. 15— 
20 thousand pelts were obtained from Russia. These figures are, 
apparently, exaggerated, for the production of the whole fur 
industry in the Russian Far East was estimated at about 40 thousand 
rubles, and hunting in these regions included sable, red fox, otter, 
Kolonok, wolverine, bear, wolf, tiger and others. Even if we suppose 
that the raccoon dog fiirs constituted 50% of this value (which is hardly 
probable), then, if the price of raccoon dog fur is 2 rubles, their number 
would not exceed 10,000. Probably, the total catch was 5—6 thousand 
pelts. 

In the 30 's, when lands were considerably opened up, about 12 
thousand raccoon dogs were caught. 

In the regions of acclimatization licensed trade started in 1948— 
1950, and the licensed restrictions were removed in 1953—1955. After 
the start of the trade, the number of animals harvested shaфly in- 
creased, and from 1953 to 1961, it fluctuated within 30—70 thousand 
pelts. For example, in 1955, about 41 thousand were obtained, and in 
1961 — 66 thousand. In the latter year, of these about 10 thousand were 
obtained from natural habitat in the Far East, and 56 thousand from 
places of acclimatization. Of these 56 thousand, Byelorussia provided 
6.5 thousand pelts, the Ukraine — ^about 5 thousand, Latvia, Lithuania 

'"According to international energy system (SI). 



121 

and Krasnodarsk territory — 4 thousand each, Kalininsk district — Ъ.1 
thousand, Pskovsk — 2.7 thousand, Astrakhansk— 2.3 thousand, 
Vologodsk, Moskovsk, Leningradsk, Novgorodsk, Smolensk, and 
Yaroslavsk districts, Azerbaidzhan, Estonia, and Dagestan — ^from 1 to 
2 thousand pelts each. In the remaining districts, territories and repub- 
lics, the annual catch was less than one thousand pelts. 

Everywhere the raccoon dog is hunted from November until the 
snow is deep, i.e. usually until February. In the Far East, the main way 
of hunting the raccoon dog is by tracking with laikas or mongrel dogs 
at night. In the past century, the Gol'dy* and Orocheny* also hunted 
by night with dogs on whose necks were fastened small bells. Less 
frequently, they hunted with different "devices" (Nadarov, 1888 and 
others). In places of acclimatization, methods of hunting raccoon dogs 
are also simple. In the majority of cases, it is hunted incidentally while 
hunting other fiir animals or specifically harvested with the help of 
dogs. With dogs, 80-90% of the animals, on average, are caught; with 
95 guns, 8-10%; with traps, 5-7%. The dog quickly follows the trail and 
overtakes the animal, and if it does not flee into a burrow, the dog 
strangles it or worries it until the hunter arrives. Traps for the raccoon 
dogs are put at burrows, along the shores of water bodies, and around 
marshes and ponds. 

Cage rearing of raccoon dogs started in the Far East in 1928 
(Firsov, 1929). In 1934 there were 15 State farms where the animal 
was raised. Collective fur farms in the first year of their existence 
selected the raccoon dog as their principal object. 

Later, the captive population in the collective fiir farms began to 
decrease because the raccoon dog requires nearly the same food as 
needed by the silver fox. By the end of the 30' s and the beginning of 
the 40's , the breeding of raccoon dog was broadly developed only in 
the collective farms of the Ukraine and from 1945 on, this type of fur 
husbandry started to decrease (Il'ina, 1952 and others). 

In the 30's work began on acclimatizing this animal in the Euro- 
pean part of USSR, in the Caucasus, and in Siberia. From 1928 to 1958 
about 10 thousand individuals were introduced in 76 districts, territo- 
ries and republics. The first attempts at transplantation started in the 
Primor'e territory where the raccoon dog was brought to some islands 
in the Japanese sea. In 1934, it was introduced into the Altai, northern 
Caucasus, in Armenia, Kirgizia, Tatariya, Kalininsk, Penzensk and 

* Indigenous tribes in the Far East — Sci. Ed. 



122 

Orenburgsk districts. In the following year — into Leningradsk, 
Murmansk, and Novosibirsk districts, Bashkiriya and other regions. 
The raccoon dog was most intensively transplanted in 1936 when more 
than 1,200 beasts were introduced, and then from 1948 to 1953, when 
500-800 individuals were introduced every year in tens of districts 
(Shaposhnikov, 1956; Lavrov, 1957 and others). 

The acclimatization of raccoon dogs in Irkutsk and Novosibirsk 
districts, in Trans-Baikaliya and the Altai did not give results due to 
severe winters and inadequate diet. The raccoon dogs acclimatized 
poorly in the montane regions of the Caucasus, Trans-Caucasus, Middle 
Asia and in Moldavia (Meladze, 1947; Yanushevich, 1956; Arens, 
1957; Trushchalova, 1959 and others). Success followed the introduc- 
tion of raccoon dog into the European part of USSR, especially in the 
northwest (Pribaltic, Byelorussia, Kalininsk, Novgorodsk, Pskovsk, 
Smolensk districts), in the central regions (Moskovsk, Yaroslavsk, 
Vologodsk, Gorkovsk, Vladimirsk, Ryazansk and other districts), as 
well as in the chernozem [black soil] belt (Voronezhsk, Tambovsk, 
Kursk districts), the lower Volga region and the level parts of the 
northern Caucasus and Dagestan. In the Ukraine, the greatest number 
of raccoon dog was established in Poltavsk, Khersonsk and Lugansk 
districts. 

The trade quality of fiir of the raccoon dog acclimatized in the 
northwest was somewhat improved. Thus, in Kalininsk district (Morozov, 
1955; Sorokin, 1956) raccoon dog fur became somewhat denser and 
softer. The length of the guard and top hairs increased by 7.96% on the 
average, that of the under fur, by 5.3%. The thickness of the guard and 
top hair decreased by 3.41%. The density of the fur increased by 
11.3%. The fi^r color became somewhat darker; the darkest "black- 
brown" color in Kalininsk district is found in 8% of the beasts against 
3% in their homeland* (Morozov, 1951, 1955; Tserevitinov, 1953). 

The importance of the raccoon dog in the hunting economy of a 
series of regions is not yet sufficiently clear. Thus, the importance of 
this animal is not clear in the Far East. The raccoon dog causes appre- 
ciable damage to the pheasant by destroying its nests; this is hardly 
compensated by the benefit gained from its destruction of harmful 
rodents and insects. In regions of acclimatization the harmful activity 
of the raccoon dog in the hunting economy, agriculture and also public 
health is obvious. Cases of nest destruction and eating of birds and 

*Probably refers to indigenous animals in the Far East — Sci. Ed. 



123 

nestlings of waterfowl and marsh birds are numerous (Komeev, 1954). 
The harm caused by the raccoon dog in floodlands and shores of es- 
tuaries is especially sensitive, since, in spring, it feeds entirely on the 
96 eggs and nestlings of waterfowl. In the floodlands of the Oka river, 
raccoon dogs destroy a great number of wild fowl; about 46% of its 
food consists of birds (Borodin, 1951). In the Voronezh preserve, birds 
constitute 48.6% of the food of the raccoon dog litters inhabiting the 
floodlands. Moreover, the raccoon dog destroys a considerable number 
of birds (45%) during the time of spring migration (Ivanov, 1962). In 
Lithuania, birds constitute 1 5-20% of raccoon dog food and this car- 
nivore causes noticeable damage to waterfowl and forest game (Prusaite, 
1960, 1961). The raccoon dog frequently destroys muskrat houses and 
eats their young (Komeev, 1954; Kritskaya, 1961). 

In the Ukraine and in many other southern regions, the raccoon dog 
causes harm to kitchen garden and melon cultivation, vineyards and 
com seedlings (Komeev, 1954, and others). Evidently the raccoon dog 
plays a negative role as a carrier of rabies, piroplasmosis and scabies 
(see above). 

A series of authors (Obtemperanskii, 1956; Yu. Popov, 1956; 
Sorokin, 1957; Samusenko and Golodushko, 1961), on the basis of the 
fact that rodents and insects constitute the main food of raccoon dog 
while birds are of secondary importance in its food, consider the harm 
caused by these beasts to be negligible. However, the destruction of up 
to 90% of the broods of waterfowl (Komeev, 1954) or of a consider- 
able number of the broods of forest game and small passerine birds is 
hardly compensated by the destruction of mouse-like rodents in the 
warm period of year. Undoubtedly, in a series of regions, the harm 
caused by the raccoon dog may be relatively small, and in others very 
great, and this requires concrete analysis; however, as a whole, damage 
by the raccoon dog to the hunting economy is evident. 

In spite of the fact that as a result of acclimatization, the prepara- 
tion of raccoon dog pelts grew 4—6 times and in a series of districts and 
republics its specific contribution to fiir production constitutes up to 
10-30%, the interests of the protection of nature and sport hunting 
economy require the destruction of raccoon dog in places where it has 
become acclimatized (A.B.). 



124 

GENUS OF WOLVES 
Genus Canis Linnaeus, 1758 

1758. Canis. Linnaeus. Syst. Naturae. Ed. X, I, p. 38. Canis familiaris 

= Canis lupus L. (domestic dog)'. 
1816. Thos. Oken. Lehrb. d. Naturg., 3, Th. 2, p. 1037. Thos vulgaris 

Oken = Canis aureus Linnaeus. 
1816. Lupus. Oken. Ibid., p. 1039. Canis lupus Linnaeus. 
1857. Vulpicanis. Blainville. Ann. Sc. Nat. Paris, Zool., 8, pt. 2, p. 

279. Canis aureus Linnaeus. 
1839. Sacalius. H. Smith. Jardine's Naturalists Library, Mamm., 25, 

p. 214. Sacalius aureus = Canis aureus Linnaeus. 
1841. Oxygous. Hodgson. Calcutta Joum. Nat. Hist., 2, p. 213. Canis 

aureus Linnaeus. 
Large species, but in part of moderate size. 
Skull broad and massive, with moderately diverging zygomatic 
arches, high (height in the occipital region more than 1/3 of condylobasal 
length), in adults and old animals well-developed crests, in particular, 
a large sagittal crest. Facial parts of the skull relatively weakly elon- 
gated, massive, quite high, and always longer than the cranial parts 
(distance from the posterior margin of the infraorbital foramen to the 
97 posterior margin of the canine alveolus is less than or equal to the skull 
width above the canines; distance from the line joining the tips of the 
supraorbital processes to the anterior end of the premaxillae is greater 
than that to the upper part of the occipital condyle). Interorbital (fron- 
tal) region convex with large cavities (sinuses) and is clearly and quite 
abruptly elevated above the line of the dorsal profile of rostrum. Su- 
praorbital processes, which also have cavities, are not large in area, but 
massive, and their upper surface is convex, the tip summit bent some- 
what downwards, and without зЬаф borders (edges). Posterior margin 
of tooth row extends to the level of the anterior margin of the 
inteфterygoid fossa. Deep notch on posterior lower margin of lower 
jaw anterior to angular process absent. 

3 14 2 
Dental formula I— C-P — M— = 42, Teeth large and massive, 
3 14 3 ^ 

'As was shown (see vol. I, p. 540), in this book domestic forms are excluded from 
the nomenclature. The wolf, C. lupus, must therefore be considered the generic type, not 
only as the most typical species of the genus but also as the ancestral form of "C. 
familiaris.'' 



125 

camassial teeth strongly manifested, palatal length about 2.5 times 
length of upper camassial tooth (P'^) and cheek teeth. Canines slightly 
curved, massive, broad at base and relatively short. When jaws are 
closed, tips of upper canines do not reach middle of mandible, while 
tips of lower [canines] do not, or only just, reach margins of alveoli of 
upper canines. Tips of upper canines directed straight downwards; no 
pronounced longitudinal sharp ridge on posterior side of canine, surface 
of canine smooth. Cutting edges of incisors with small accessory lateral 
cusps (trilobed). 

In general appearance, the animals are long-legged, with notice- 
ably elongated trunk; tail is fluffy, of moderate length — up to two-thirds 
length of trunk with the head (often less) and not more than twice as 
long as hind limbs. When the animal is standing, its end is usually not 
lower or only slightly lower than the tarsal joint, but it never reaches 
the ground. 

Head notably elongated with somewhat pointed muzzle or else 
blunter or fairly broad with broad forehead. Ears relatively short with 
pointed tips. On sole of hind foot, behind four naked digital pads is 
situated a larger fifth. Pads of digits III and IV of fore limbs either 
separate or united (jackal, Fig. 17). 

Fur quite coarse. Color unimoфhic (dimoфhic only in some races 
of wolf, C. lupus, fi-om southern North America — ^"C. niger"), grayish, 
yellowish, and reddish tones mixed with black hairs, sometimes with 
noticeable black dorsal saddle; adult animals never exclusively mono- 
tone (except black phase of "C. niger"). Violaceous* gland present, 
but weakly developed. Teats usually 5 pairs. 

Sharp sexual dimoфhism absent, but females are usually some- 
what smaller than males. Age dimoфhism occurs, and seasonal 
[dimoфhism] is well marked in some species, but manifested almost 
only in density and length of ftir. Two molts per year. 

Species of the genus are very close to each other and homogeneous, 
representing the wolf type— the true gray wolf (C. lupus) and its small 
copy — the jackal. The large forms of the wolf have the greatest dimensions 
(see family characteristics, and below — description of wolf), the smallest 
jackals are not more than 80-90 cm in body length, usually about 70-75 
cm, and in weight not more than 10-12 kg, usually about 6-7 kg. 

Species of the genus are found in all landscapes from tundra to 
lowlands of hot deserts, and high mountains. They prefer regions with 

*Also called supracaudal — Sci. Ed. 



127 

more or less open areas and abundant wild or domestic hoofed mam- 
mals. They are sedentary, [but] individual animals sometimes roam 
widely and settle in distant areas. Monogamous animals; moreover, 
some form close pairs for several years. 10—13 pups, usually less. 
Young usually delivered in open den. Predators, feeding mainly on the 
meat of mammals, frequently large ones (ungulates), in part on carrion 
99 and other vertebrates, exceptionally on invertebrates, and at times even 
plants. Except during breeding, they usually live and hunt in small 
groups. In some species, internal relationships in the group (rank, etc.) 
are shaфly marked (wolves), in others, quite weakly (jackals). 

This genus is very widely distributed, extending in the north to the 
northemmost points of land (83°20' N. Lat. — not found on floating ice); 
also found in equatorial regions. In the New World, it occupies the 
entire continent of North America (including the entire Arctic archi- 
pelago) and in the south, it extends to Costa Rica inclusive. It is absent 
on the Queen Charlotte Islands and islands of the Bering Sea. In 
Greenland, it occupies the northern, eastern and western coasts, south- 
ward nearly to 70° N. Lat. 

In the Old World, its range occupies all of Africa (excluding Mada- 
gascar) and all of Eurasia southward to southem China, Siam [Thailand], 
Burma, Assam and Ceylon inclusive. In the north, the range includes the 
Kolguev Islands [Barents Sea], south island of Novaya Zemlya, Belyi 
[island] adjacent to the Taimyr coast, as well as Lyakhovskii Islands and 
the New Siberian archipelago. They are also found in Kamchatka, Sakhalin 
and the Japanese islands. They are absent on Iceland, Spitsbergen, Franz 
Josef Land, Sevemaya Zemlya, Wrangel Island, the Commander Islands, 
the majority of the Kuril islands and Shantar Island, Taiwan, Hainan, and 
the Malacca and Malaya archipelago.^ At the present time, it is extermi- 
nated in a series of territories of North America, in the greater part of West 
and Central Europe, in part of Japan and in several other places. This genus 
represents a highly adaptable and viable group, and its range has decreased 
only in the regions of dense human population and where it has been 
directly persecuted to extinction. 

The genus appeared in the upper Pliocene of North America; in 
Eurasia, in the Pleistocene (see characteristics of genus Vulpes). 

Particular generic or subgeneric names have been proposed for 
almost every species included in the genus of contents accepted here. 
This occurred during the period of narrow inteфretation of the species 

^A feral domestic dog — ^"Canis dingo" — is found in a completely wild condition in 
Australia (except Tasmania). 



128 

concept, when many of the races were considered to be separate spe- 
cies, and these [genera or subgenera] could be group names. At the 
present time, there is no basis for dividing the genus into several 
subgenera, and it is senseless to separate, within it, such subgenera as 
Canis for the wolf and Thos, Schaeffia, Lupulella for the various forms 
of jackals. The differences between these "subgenera" are very insig- 
nificant; moreover, in the present species concept, only one species is 
included in each subgenus. Division of the genus into two subgenera: 
Canis and Thos, i.e. wolf and jackal is also not justified. In some 
respects, the species of the genus themselves represent the fullest and 
most characteristic expression of the family type. 

This genus is most closely related to the group Simenia (Abyssinian 
wolf) and Vulpes (foxes) and especially to Alopex (arctic fox). This 
latter represents, in craniological respects, a quite clear transition and 
connecting link between Canis and Vulpes. In the past, all these groups, 
especially Canis, Vulpes and Alopex, were usually grouped in one genus 
or were considered as subgenera only. Later, however, all authors es- 
sentially divided these groups generically or, at least, separated Canis 
from Vulpes, united Alopex with the latter group, and separated Simenia. 
Most recently, it was again proposed (Haltenorth, 1958) that all the 
mentioned groups including a series of others be considered members of 
one enlarged genus — Canis (see characteristics of the family). In this 
account, the described groups occupy subgeneric positions. This point of 
view deserves consideration, but requires broader argument. 

In the genus are 5 species: 2 African — C. adustus (striped jackal) 
and С mesomelas (shabrack [= black-backed] jackal), 1 North Ameri- 
can — C. latrans (coyote), 1 Eurasiatic— African — С aureus (jackal) 
100 and 1 Eurasiatic— North American — C. lupus (wolf).^ The species of 
this genus constitute about 1 7% of the species of the family. 

One species — the wolf— is found both in the wild and in the domestic 
state as the domestic dog CCanisfamiliaris"). The dog — the most strongly 
altered form as compared to its wild ancestor — is generally the most strongly 
divergent and diverse of domestic animals (the number of breeds exceeds 
200). Nevertheless, in respect to the wolf, there is no reproductive isolation. 
The view which was widespread not long ago, concerning the diphyletic 
origin of the domestic dog — from wolf and jackal — is now totally dis- 
regarded. As in other domestic animals, the dog has one original species 
("ancestor"). The possibility of the diphyletic origin of the dog is excluded 

^The wolf of Florida and the southeastern states adjacent to the Mississippi 
[River], is considered by American authors to be a separate species — С niger. 



129 

in particular by the difference in the diploid number of chromosomes — 78 
in wolf and dog and 74 in jackal (Matthey, 1954). 

In the USSR, two species are found: wolf, Canis lupus Linnaeus, 1758 
and jackal, Canis aureus Linnaeus 1758 — about 40% of the species of the 
genus and about 0.6% of the number of mammals of the Union. 

The range of the genus covers the entire territory of the USSR, 
except the above-mentioned northern territories and islands. 

The species of the genus are of great importance in the USSR as 
pests of livestock and the hunting economy and, to a lesser extent, as 
carriers of rabies. Their importance as fur-bearing species is not great 
and less than the harm they cause (V.H.). 

JACKAL, CHEKALKA 

Canis aureus Linnaeus, 1758 

1758. Canis aureus Linnaeus. Syst. nat. Ed. X, 1, p. 40. Laristan, 

southern Iran. 
1835. Canis aureus var. moreotica. Geoffroy. Exped. Scient. de Moree, 

Zool., pi. 1. Peloponnesus (Morea), Greece. 
1858. Canis aureus typicus oder var. caucasica. Kolenati. 

Reiseerinnerungen. Th. I, p. 96, Armenia. (V.H.). 

Diagnosis 

Adult body length not more than 90 cm, tail length about one-third of 
body length. Length of skull less than 200 mm. Arciform line formed 
by anterior border of both nasal bones has, in the middle (along suture 
between nasal bones), a well-marked projection, directed anteriorly. At 
antero-intemal sides of auditory bullae, swellings in form of ridges on 
basioccipital absent. Cingulum on outer surface of first upper molar 
well marked, broad and continuous (Figs. 15, 16). (V.H.). 

Description 

In its general appearance, the jackal is very similar to the wolf, but 
much smaller in size, lighter in weight, with shorter legs and with a 
somewhat more elongated trunk and shorter tail. Its end just reaches the 
heel or a bit below it. As in the wolf, the tail always droops. The head 



130 

is lighter, not so "foreheaded"*, and the muzzle is narrower and more 
pointed. On the whole, the jackal much resembles a small mongrel dog. 
Fur is coarse and stiff. Iris is light or dark brownish. Teats 5 pairs. 
General color of winter fur is dirty reddish-gray, strongly high- 
lighted with blackish due to the black tips of many guard hairs, or a 
brighter, rusty-reddish color. Anterior part of muzzle, circumorbital 
region and forehead are ocherous-rusty-reddish. Above each eye, a 
101 blackish stripe is present. Margins of lips and lower cheeks are dirty 
white. Upper part of forehead and occiput are ocherous. Back of ears 
is pale-rusty; ears are covered internally with dirty whitish hairs. Chin 
and throat are whitish, with a dirty tint. 

Black guard hairs are especially developed on the back, but less so 
on the sides, and general color here is brighter and clearer. Belly is 
whitish along the midline, and in the axillary region and groin the color 
is mixed with a reddish tint. Extremities are ocherous-red, with internal 
surfaces of lighter color. Tail is gray with an ocherous tint and a 
strongly defined dark color on dorsal side and at tip. General color 
intensity and the degree of development of darkening is quite strongly 
variable individually, and both comparatively dull jackals as well as 
those with very bright colors are encountered. 




Fig. 36. Jackal, Canis aureus L. Sketch by A.N. Komarov. 

•Russian word is "lobastaya", referring to the more expanded frontal region of the 
wolf compared to the jackal — Sci. Ed. 



103 



131 

Summer fiir is more sparse, coarse and short and has same color 
as that of winter, but brighter with dark tint less developed. Sexual 
differences in color are absent. Pelage of newborns is very soft and its 
color varies from light-gray to dark-brown. This pelage remains on the 
animal for about one month and then the first adult coat starts to grow 
and is developed by August. It is very similar to that of adults. Geo- 
graphic variation in color is fairly noticeable. 

The skull is similar to that of the wolf in its main features and in 
general appearance, but is smaller and less massive. The nasal region 
is lower, and the facial shorter (its length is nearly equal to that of the 
cranial region), dorsal profile of the facial region is nearly straight and 
depression in middle of nasal bones is very weak. In the region of the 
camassial teeth, the skull is not broadened and its outline, beginning 
with the line of the zygomatic arches, converges anteriorly in a quite 
regular wedge (on looking up at the skull from below, a sharp projec- 
tion in front of the camassial tooth is not seen). Sagittal and occipital 
crests are well developed, but weaker than in the wolf Supraorbital 
processes are large, but relatively less massive than in the wolf. Be- 
tween the frontals and between the nasals, along the corresponding 
sutures, a shallow longitudinal depression is present. On the line of the 
anterior margins of the nasal bones (at suture between 2 nasal bones) 
there is a distinct projection (Fig. 15). At the anterior internal borders 
of the auditory bullae at the basioccipital, the bulla is not bordered by 
bony ridges. 

Canines are large and strong, but relatively thinner than in the 
wolf, and camassial teeth are relatively weaker. The cingulum on the 
outer margin of the first upper cheek teeth is continuous, broad and 
well defined throughout its whole extent (Fig. 16). 

Sexual differences in the skull are only manifested in the generally 
somewhat smaller dimensions of the female. Age variations correspond, 
in all principal aspects, with those described in the wolf. 

Diploid number of chromosomes — 74. 

Male body length 71—85 cm (as a rare exception in the species, 
may be somewhat more), females average somewhat less."* 

'•In regions on the right tributaries of Pyandzh it was shown (Chemyshev, 1958) 
that body length of males (14) is 68-104 cm, and of females (17) is 71-100 cm. These 
maximal figures are, apparently, exaggerated. Such figures are not given by any of our 
authors for either Middle Asiatic or Caucasian animals. Mean body length, drawn from 
the tables given by the mentioned authors, is 81.5 cm for males and 80.5 cm for females. 
The maximal specimen, of 75 caught in India, Baluchistan and Iran had a body length 
of 81 cm (male) and 73.5 cm (female). Six females from Nepal had an average body 



132 

Measurements of animals from southern Tadzhikistan (region of 
the right tributaries of Pyandzh and upper Amu-Dar'ya) are as follows. 
Adult (sexually mature) males: tail length (14) 22-27 cm, length of 
hind foot (13) 13.7-19 cm, height of ear (8) 6.7-10 cm. Aduh females: 
tail length (17) 20-27.5 cm, length of hind foot (17) 9.5-12 cm, height 
of ear (15) 7.1-9.1 cm. Height at shoulder (12) of males and females 
44.5-50 cm, height at sacrum 40-48 cm. 

Weight of males (15) 6300-13,670 g, of females (16) 7000- 
1 1,240 g. 

Weight of heart 100-150 g, of liver 185-480 g, total length of 
intestine 335-454 cm, length of caecum 7.5-12 cm (Chemyshev, 1954). 

Measurements of male skulls (14 from Caucasus and from Turkestan; 
Ognev, 1931; ZMMU*): condylobasal length 147-164 mm, zygomatic 
width 79—97 mm, breadth of muzzle above canines 27—32 mm, 
interorbital 22.6-30.2 mm. Skull of females (17) from the right bank 
ofupper Amu-Dar'ya (Chemyshev, 1958): condylobasal length 123.7- 
159.7 mm, zygomatic width 69.9-92.5 mm, length ofupper toothrow 
64.5-87.5 mm; interorbital width 19.4-30.1. 

Measurements of jackals within the USSR do not show clear geo- 
graphic variation. (V.H.) 

Systematic Position 

The jackal must be considered as a somewhat less specialized form of 
the genus than the wolf. This is manifested in the relatively short facial 
portion in the somewhat weaker tooth row, with which, in particular, 
the above-mentioned "wedge-shaped" skull and its relatively small width 
in the region of camassial teeth is connected, and in the less developed 
crests. This is, apparently, connected with its diet of small prey — birds, 
rodents, lower vertebrates, insects, other animals and carrion. The jackal 

length of 73 cm, 12 females from Palanpur and Sind — 69 cm (Pocock, 1941). It is 
generally considered that Indian animals have a body length of 61-76 cm (Prater, 1947). 
Maximal Balkan jackals had a body length of 85.5 (male) and 81 cm (female; Atanasov, 
1953). Our jackals are. apparently, slightly larger than the Indian ones, but not by much. 
It is considered, under the circumstances, a mistake, that the skull dimensions given for 
Tadzhikistan, do not differ from generally known ones. One foreign author (Van den 
Brink, 1958), who did not refer to the source, gave still more exaggerated dimensions 
for the European jackal: 85-105 cm, and another (Marches [et al.], 1954) gives a body 
length of 104 cm for a male and 100 cm for a female from Romania. A weight for jackal 
of "more than 16 kg" (P. Manteifel; Chemyshev, 1958) is also exaggerated. 
*Zoological Museum, Moscow State University. 



133 




102 



Fig. 37. Skull of jackal, Canis aureus L. 



134 

is a "scavenger" in no lesser degree than it is a predator. This is, 
apparently, the significance of the somewhat smaller general dimen- 
. sions of the animal (V.H.). 

Geographic Distribution 

Southern Asia and Near East, southeastern Europe, northern half of 
Africa. 

Geographic Range in the Soviet Union 

This constitutes a considerable part of the species range — its northern 
border is connected with the southern part of the country — Caucasus, 
Middle Asia and Moldavia. It consists of several parts within the Soviet 
Union separated from each other which unite beyond our borders. 
104 In the Caucasus, the range of the jackal is connected with the 
plains and foothills, in places with low and even middle montane belts, 
and having quite complicated contour. In the western half of the coun- 
try, the jackal is distributed in a narrow strip along the entire Black Sea 
coast from Novorossiisk to Batumi. Here, the animals are not usually 
distributed in mountains higher than 400-500 m above sea level. The 
range widens somewhat along the Rion Basin and extends somewhat to 
the east of Kutais'. 

In the beginning and middle of the past century, the jackal was, 
apparently, distributed (perhaps in part as vagrants, however this was 
not very rare) along the eastern shore of the Azov Sea to the lower Don 
and even to Taganrog. Old reports about the occurrence of the de- 
scribed species in Crimea are not worthy of attention. 

In the eastern half of the Caucasus, beyond the Main Range, the 
jackal is distributed on the plains and foothills and, as a rule, is not 
found on mountains higher than 800-900 m above sea level. Only in 
Zakatal, in autumn, it may climb up to 1000 m searching for fruits, and 
everywhere in Talysh, to a height of 1700 m (the highest occurrence 
known in the USSR). In the Trans-Caucasus, the jackal is distributed 
from Talysh and the Lenkoran lowland southward to the Apsheron 
peninsula and the foothills of the Great Caucasus on the north. Here, 
it extends to Zakatal and a little more to the north. In the west, the 
jackal is distributed along the lowland places of the lora, Alazan' and 
Kura basins, to the Tbilisi meridian and a little to the west (Mtskhet). 
Southward, the foothills of the Little Caucasus constitute the western 
border of the eastern Trans-Caucasian part of the range. 



135 

Along the valleys of various rivers, the jackal penetrates in some 
places in a minor way into the mountains. Along the Araks valley, the 
range extends in a narrow strip westward to Oktemberyansk region 
(44° E. Long., a little east of Erevan). In the Araks, the animal does 
not ascend higher than 500-850 m above sea level. 

From the Apsheron peninsula, the range of jackal extends, in a 
narrow strip, along the Caspian Sea northward to the mouth of the 
Terek and Kuma and to the Beryuzyak peninsula. In part of the range 
northward to Makhachkala, the jackal is distributed along the sea shore, 
generally not further than 20—30 km from it. Beginning only at 
Makhachkala, it goes deeper into the country along the foothills and 
river valleys. To the west of Makhachkala, the jackal is distributed in 
a narrow strip along the foothill belt to Khasav"yurt and a little west- 
ward. It goes west along the Terek to Shelkova (Shelkozavodsk) and 
Parabochevsk forest and reaches Mozdok, and along the Sunzha, to 
Groznov. It is not found along the Kuma river upstream from its mouth. 

In the past, jackals (apparently, mainly along the Caspian coast), 
reached the delta of the Volga (30's and 40's of the nineteenth century) 
and were also found in "Don steppes" and still in the middle of the past 
century along the Ergeni river. They also inhabited Stavropol' at that 
time. It is now impossible to decide whether the places of occurrence 
along the Ergeni and Don were connected with the Pri-Caspian region 
of occurrence of the species, or with the Pri-Azov region. It is probable 
that all these were one region. 

Of the Caspian Sea islands, the jackal lives only on Sar Island at 
Lenkoran'. It is absent on other even larger (Chechen') islands as well 
as on Agrakhansk peninsula (spit). 

In the last ten years, to a considerable degree in connection with 
the drying of the Caspian Sea and the disappearance of reed beds, the 
range of the jackal in the northern Caucasus has contracted. In particu- 
lar, it does not reach Kuma. 

In the northern Caucasus, chiefly at the beginning of the present 
century, fairly long intrusions are known — ^to Gunib in the heart of the 
mountains of Dagestan, to Stavropol', to Psebai (south of Armavir), to 
the mouth of the Kuban' river and even into the mouth of the Don. 
Wanderers in the eastern parts apparently came from the Kizlyar re- 
gion of occurrence of the animal, and in the west, from the Black Sea 
coast. Jackal intrusion even to Tambov (Ognev, 1931) is known, prob- 
ably from the Caucasian region of occurrence of the animal. 
106 The distribution areas of the jackal in the western and eastern 

halves of the Caucasus are, in essence, separated from each other, but 



137 

in the middle of the Trans-Caucasus they are very near to each other, 
separated only by the Suramsk range and the adjacent regions — a total 
of 100 or 100+ km. Animals occur only as intruders and their move- 
ments, apparently, reflect some kind of connection that is or was realized 
between the two described divisions of the range. The present union of 
the above-mentioned parts of the distribution area applies, however, 
only farther the south, outside the boundaries of the USSR.^ 

The Middle Asiatic region of jackal occurrence within the Soviet 
Union is completely isolated from that of the Caucasus. It has quite a 
complicated form. This is explained by the fact that the jackal, in its 
distribution here, is associated with river valleys and cultivated lands, 
resolutely avoids deserts, and does not go high into the mountains. It 
is, in part, divided into separated sections, isolated from each other. In 
the west, in Turkmeniya, the jackal is found along the Atrek, from 
Chikishlyar westward to the mountains. Along the river valleys of the 
Atrek, Sumbar and Chandyr, it penetrates into the western parts of the 
Kopet-Dag (at least to Kara-Kala on the Sumbar and Yarty-Kala on 
the Chandyr), but it does not get into the heart of the Kopet-Dag 
mountains. Its range envelops the Kopet-Dag from the west (?) and 
north as a narrow strip along the foothill plains and foothills, and 
extends to Tedzhen^. The jackal occupies the entire Tedzhen valley to 
its extreme lower reaches and all the valley of the Murgab with its 
upper tributaries (Kushka, perhaps Egri-Gek). In the desert between the 
Tedzhen and Murgab, and between the Murgab and Amu-Dar'ya, the 
jackal is absent. The union of these sections is realized in Iran and 
Afghanistan. 

Along the Amu-Dar'ya, the jackal is distributed from the Chubek 
(on the river, somewhat east of the mouth of the Kizyl-su, nearly 
directly south of Kulyab) to the delta which occupies the whole region. 
In the basin of the upper Amu-Dar'ya, the jackal is distributed along 
the Kizyl-su and Yakh-su northward, nearly to the latitude of Kulyab 
or somewhat farther north; along the Vakhsh, it goes up to Kurgan- 
Tyub; along the Kafirnigan, nearly to Dushanbe or a little to 
the northeast, to the mouth of the Varzob; along the Surkhan-Dar'ya 

^The distribution in the Caucasus according to materials of Nordmann, 1840; 
Komilov, 1859; V.V. Shch., 1887; Alferaki, 1910; Satunin, 1915; Ognev, 1931; Heptner 
and Formozov, 1941; Dal', 1954; Heptner, Turova and Tsalkin, 1950; Kirikov, 1959 and 
the unpublished material of V.G. Heptner. 

^The distribution of the jackal in this region is poorly delimited. Apparently, the 
mentioned strip is not continuous. In sand and loess deserts, the jackal is absent. It is 
also absent in the Great Balkhan and hardly reaches the West Uzboi. 



138 

upstream, at least to Regar. In this way, the range includes the 
entire Gissar valley and the foothills of Gissar range eastward to 
Ordzhonikidzeabad and almost to Ramit. Between the Surkhan-Dar'ya 
[river] and Kugitangtau and Baisun[tau] mountains, the range protrudes 
northward from the Amu-Dar'ya, extending to the city of Chirabad. In 
all the described sections on the right bank of the Amu-Dar'ya, the 
jackal does not ascend higher than 1000-1 100 m above sea level and 
the boundary of its distribution envelops, as it were, the higher regions 
such as Babadag and other massifs lying along watersheds, and forms 
here a fairly complicated pattern. 




107 Fig. 39. Reconstructed species range of the jackal, Canis aureus L. V.G. Heptner. 



139 

The range of the jackal further occupies the entire area of culti- 
vated land along the Kashka-Dar'ya (Karski, Chakhrizyabs). This region, 
apparently, is more or less isolated from the previously mentioned 
Baisuntau mountains. The jackal exists everywhere in the Zeravshan 
basin (Bukhara and Samarkand oases), penetrating upwards along the 
river to the mountains, to Pendzhikent, and perhaps somewhat higher. 
The jackal inhabits the cultivated strip and adjacent places between 
Samarkand and Tashkent and is widely distributed in Tashkent oasis, 
eastward to the mountains. Along the Syr-Dar'ya, it penetrates to the 
western parts of the Fergana valley and to the Farkhandsk reservoir and 
Kanibadam (for the eastem parts of the valley, almost nothing is known). 
Downstream along the Syr-Dar'ya, the jackal is normally distributed 

107 to the mouth of the Aris. East of the Syr-Dar'ya, at the northern limits 
of its distribution, it extends to Shimkent, Lenger, Georgievk' and 
Boroldaisk mountain, and to the sources of Bugun river lying approxi- 
mately 150 km north of Chimkent. 

In Middle Asia, distant intrusions of the jackal are also known and 
moreover they take place partly in years of severe cattle losses (con- 
tinuous snow cover)*. In these years, intrusions are more frequent and 
extend particularly far. Intrusions of jackals may occur in the lower 
Syr-Dar'ya, reaching Kazalinsk; on the Sary-su river; on the Chu river 
to its lower reaches; to the northwestern edge of the Betpak-Dala desert 
and almost reaching even to the city of Karaganda. To the north intru- 
sions are known to the Ustyurt (Kos-Bulak mountain), on the 
northwestern shore of the Aral Sea (Kintekchi), in the Asmantai-Matai 
sands at the northern base of the Ustyurt, at Emba. Jackals wandered 
Turgai and Sary-Kopa and even to Orenburg (see below for details)^. 

The third region in which the jackal is known in the USSR is 
Moldavia. Here, the animals are rare, appearing as vagrants, and not 
coming this far every year, in the southern and southwestern regions. 

108 They enter from Romania (Kuznetsov, 1952). The appearance of the 
vagrant jackals in the Trans-Carpathian region cannot be excluded. 



*The Mongolian word dzhut refers to persistent continuous snow cover that pre- 
vents ungulates from feeding — Sci. Ed. 

^Range in Middle Asia according to Ognev, 1 93 1 ; Laptev, 1 934; Flerov and Gromov, 
1934; Flerov, 1935; Shestoperov, 1936, 1936a, 1936b; Gromov, 1937; Leviev, 1939; 
Salikhbaev, 1939; Sludskii, 1939; Sultanov, 1939; Kuznetsov, 1948; Bazhanov, 1951; 
Shukurov, 1951; Afanas'ev et al., 1953; Chernyshev, 1954, 1958; Heptner, 1956 and 
others and according to the unpublished material of V.G. Heptner. 



140 

Geographic Range outside the Soviet Union 

The range in Asia includes southwest Siam, Burma, Assam, Bengal, all 
of India; in the north, it includes Bhutan, Sikkim, Nepal (in Himalaya 
to a height over 3000 m above sea level), Ceylon, Baluchistan and 
Afghanistan (except the montane areas of the Hindu Kush), Iran, Asia 
Minor and the entire Arabian peninsula. In Africa, the range includes 
Kenya, Ethiopia, Somalia, Sudan, Egypt, Libya, Tunisia, Algeria, 
Morocco, Rio de Oro [Spanish West Africa, now Western Sahara], and 
Senegal. In Europe, the jackal is found in Greece, Albania, Yugoslavia 
and Bulgaria. As a vagrant known in Hungary (in the north to Komitata 
KJievesh near Tissa northeast of Budapest) and in Romania (very rare, 
only in the southwestern part of the country at the Danube south of 
Kraiov)l (V.H.). 

Geographic Variation 

Throughout a broad area covering the country, with extremely different 
natural conditions, geographic variation of the jackal is quite well 
defined, though less than what is accepted. Features of size as well as 
color vary. In the USSR, there occur two well-differentiated forms. 

1 — ^Turkestan jackal, C. a. aureus Linnaeus, 1758. 

General color is relatively pale with sandy tones predominating. 
The fur is relatively soft. Dimensions large. 

Middle Asia. Outside the USSR — ^Afghanistan, Iran, Iraq, Arabian 
peninsula, Baluchistan, northwestern India (Kutch, Sind, Gujarat). That 
animals inhabiting southwestern Turkmeniya are not completely iden- 
tical to those living east of the Amu-Dar'ya is not excluded. 

2 — Caucasian jackal, C. a. moreotica Geoffroy, 1835 (syn. typicus, 
caucasicus). 

General color is bright and saturated, with strongly pronounced dark 
blackish tones on the back. Thighs and upper parts of legs, ears and 
forehead are bright, reddish chestnut. Fur is coarse. Size somewhat less. 

Caucasus, Moldavia. Outside the USSR — Southeastern Europe, Asia 
Minor. 



''Calinescu. 1930; Atanasov, 1953. Judging by the catch in Moldavia, one may 
consider that intrusions occur also in other parts of Romania and take place more 
frequently than is known. 



141 

In territories lying beyond our borders, the following forms are usually 
accepted: C. a. syriacus Hempr. et Ehrenb., 1833 (Syria, Palestine); С 
a. lupaster Hempr. et Ehrenb., 1833 (Libya, Egypt, Palestine); C. a. 
indicus Hodgs., 1833 (Nepal, Sikkim, Bhutan, Assam, Burma, Siam); 
C. a. naria Wrought., 1916 (south part of India); C. a. lanka Wrought., 
1916 (Ceylon); C. a. algiriensis Wagn., 1841 (Tripoli, Algeria, Tunisia); 
С a. maroccanus Cabrera, 1921 (Morocco); С a. soudanicus Thos., 
1903 (Egypt, Nubian desert, Kordofan, Sudan). Some of these forms 
are highly doubtful. Sometimes the accepted (Ellerman and Morrison- 
Scott, 1951) C. a. cruesemanni Matschie, 1900, described according 
to a living specimen by an author of very little reliability, is not taken 
into account here. The existence of the name ecsedensis Kretzoi, 1947 
(hungaricus Ehlik, minor Mojsis.) given to animals from Hungary is 
also not credited. These are, undoubtedly, vagrant specimen of the 
Balkan moreoticus; jackals do not live permanently in Hungary. 

The statements that there are 20 species of jackals in the Old 
World, not including the black-backed (Ognev, 1931), is completely 
unsubstantiated. The true jackals, together with the black-backed, are 
only 3 species {aureus, adustus, mesomelas). The previous conception 
was based on binary designation of races, the unnecessary division of 
species and subspecies and upon unreviewed synonymy. (V.H.) 

109 Biology 

Population. Along the Caucasian coast of the Black Sea, at Novorossiisk, 
the jackal is rare; to the south, it is common, while in Georgia it is 
abundant. At the mouth of Terek, it is rare; southward, its number 
increases, reaching a maximum in southern Azerbaidzhan. In Dagestan, 
from 1936 to 1941, an average of 580 jackal furs were prepared yearly. 
The jackal is abundant in the Kolkhid, Zakatalo-Ismailinsk, Kurinsk, 
Lenkoran and other lowlands in Georgia and Azerbaidzhan. In Arme- 
nia and Nakhichevan Autonomous SSR, they are encountered in small 
numbers. In the Trans-Caucasus, in the 30's of the present century, up 
to 17 thousand jackals were caught yearly. In the Trans-Caucasus, 
jackals are most common in Azerbaidzhan. From 1931 to 1940 an 
average of 9400 animals was caught here yearly, in Georgia — ^2300, in 
Armenia — some hundreds (Vereshchagin, 1947). 

In Turkmeniya, it is abundant in the valleys of rivers and rare in 
the foothills and low mountains, from whence, from time to time, it 
penetrates in small numbers to the very edge of the desert. In higher 



142 

areas deep within the Kopet-Dag, it is absent or rare (V.G. Heptner). 
On the lower Atrek, it is not numerous and comparatively few are 
captured (106 in 1941; Samorodov, 1953). It is rare in West Uzboi. In 
the valleys of the Tedzhen, Murgab, and Amu-Dar'ya rivers, and in the 
oasis irrigated by them, it is abundant; and is more rarely met with 
along the Sumbar and Kushka. In the mountains of southern Turkmeniya, 
it is encountered regularly. It is known in Karabil', the mountain be- 
tween the Kushka and Murgab rivers, and on the Gyaz'-Gyadyk (upper 
Tedzhen), but in small numbers (V.G. Heptner). From 1948 to 1958, 
from 1400 to 5400 jackals were caught annually in Turkmeniya 
(Sapozhenkov, 1960), 

In Tadzhikistan, jackals are abundant in the valleys of the Pyandzh, 
Vakhsh, and Amu-Dar'ya rivers and their tributaries — the Kafimigan, 
Kyzyl-su, Surkhan-Dar'ya and in the adjacent cultivated lands. In the 
offshoots and foothills of the mountains they are rare. It is especially 
numerous in the Pyandzh tugais* from Faizabad-Kala to the mouth of 
the Vakhsh and along the entire Vakhsh valley where, however, they 
avoid sandy deserts. In the settlement of Tigrov Balka in the Pyandzh 
valley, in 1946-47, on an area of 600 hectares, about 133-161 jackals 
were counted, i.e. up to 0.02 individual per hectare. In the mountains of 
the Vakhsh-Kafimigan interfluve, it is encountered only during migra- 
tions. To the west of the Kafimigan, and to the north of the Amu-Dar'ya 
tugais, it is very rare in the foothills of Koikutau and Tuyuntau, inhab- 
iting the whole of the cultivated strip of the Gissar valley, and it is also 
not rare in the foothills of Gissar range (Flerov, 1935; Chemyshev, 
1954, 1958). It is common in the valleys of the upper reaches of the 
Syr-Dar'ya and Zeravshan rivers (Chemyshev, 1954). From 1951 to 
1958 from 530 to 1400 jackals were caught in Tadzhikistan. 

In Uzbekistan, the jackal is numerous in the valleys of rivers 
Surkhan-Dar'ya, Kashka-Dar'ya, Zeravshan, Syr-Dar'ya, in the lower 
Amu-Dar'ya and in Khorezm and Bukhara oases; it is rare in foothills 
and also in the Fergana valley in the floodplains of the Naryn and Syr- 
Dar'ya rivers. In Fergana, it is captured occasionally (Malenkov, 1958). 
Only in the lower Amu-Dar'ya, in the territory of Karakalpak Autono- 
mous SSR, are about 1000 jackal furs prepared yearly (Kostin, 1956). 
In the Amu-Dar'ya delta about 200 jackals were estimated in 1948 on 
six tugai woodlots (Salikhbaev, 1950). From 1950 to 1958, from 3300 
to 6200 individuals were prepared annually in Uzbekistan. 

*The Russian tugai refers to a distinctive riparian shrubland found along rivers of 
Middle Asia — Sci. Ed. 



143 

It is rare in the Syr-Dar'ya valley in its middle course north of 
Kzyl-Orda, but in some years, it is very common from its confluence 
with the Arys' and farther south, and also in the spurs and foothills of 
the Talask Alatau in the territory of Chimkentsk district. 

Habitat. In the Caucasus and Trans-Caucasus, the jackal is a low- 
land dweller, and as a rule does not ascend into the mountains higher 
than 600 m above sea level. Only in a few places (Borzhomi), it is 
found at a height of 900-1050 m (Dinnik, 1914), and in Armenia at 
present climbs to heights of 840 m (Dal', 1954). It inhabits the coasts 
of the Black and Caspian seas and the level valleys of rivers every- 
10 where where there is dense forest, thickets of prickly bushes and 
extensive reed floodlands rich in game and small rodents. Where dense 
shrubs are absent, jackals are absent. Preferred habitats are impassable 
thickets of bushes, consisting of various spiny and climbing plants such 
as blackberry, Jerusalem thorn {Paliurus australis), Smilax excelsa. 
Clematis sp. and others. In these thickets, the yearly falling leaves and 
the dry branches form, at some height from the ground, a unique floor- 
ing that divides the thicket into two levels: in the lower, the inhabitants 
of these thickets move freely — jackals, jungle cats, badgers, wild boars, 
pheasants and others feeling themselves here to be in comparative 
safety (Satunin, 1915). Such habitats serve as shelters for jackals in 
southern Dagestan, on the lower Samur. They are nearly inaccessible 
to humans, but the animals freely move there along "tunnels" made by 
wild boars (Heptner and Formozov, 1941). 

The jackal hunts along rivers and canals, on the sea shore and 
around lakes, beating down, together with other animals, the trail sys- 
tem. In dry years, when lakes and swamps dry out strongly, jackals 
concentrate around them, hunting waterfowl, nutria and small rodents. 
In severe winters, when water bodies are covered with ice, the carni- 
vores gather around them, cross over to the island and settle in the 
reedbeds away from the shore and through the course of the entire 
period of frost, do not leave the water bodies. In spring, summer and 
autumn, 70 jackals and 60 jungle cats were caught in a small section 
of the Inkitsk swamps (Georgia) (M. Pavlov, 1953). Jackals live very 
close to settlements. One time a jackal lived in the center of Lenkoran 
village under a forester's house. 

In Middle Asia and Kazakhstan, the jackal is also a lowland dweller. 
Here, the preferred habitats are tugai thickets and reed floodlands in the 
floodplains of rivers, bushes and reeds along the canals and aryks* and 

*The Russian atyk is a Turicic loan word for irrigation ditch — Sci. Ed. 



144 







Fig. 40. Habitat of jackal. Outer margin of tugai of tamarisk and turanga, in 
the Murgab valley at Imam-baba, Turkmeniya, 22 May 1962. Photograph by 

A.A. Sludskii. 

also thickets on abandoned irrigated lands. They prefer wild olive- 
turanga and tall grass-herb tugais. Here, trees are densely interwoven 
with vines and surrounded with sturdy thickets of reeds or plume grass 
[Erianthus]. Places where jackals live usually abound in waterfowl, 
pheasants, tolai hares, small rodents, frogs and fish. Here it finds good 
cover and abundant prey. 

In the Gissar and Fergana valleys, jackals do not find hiding places 
in the arable fields in the valley, and live in adjacent low hillocks 
[called] adyr. In the adyrs it shelters in dry channels, loess caves and 
niches, occupying abandoned holes of foxes and porcupines, or digging 
its own holes. In Turkmeniya, if there arenot good thickets along the 
river, as for example on the Kushka, the jackal also makes a hole found 
in the neighboring hillocks (Heptner, 1956). 

The jackal willingly settles in oases, where it confines itself to 
thickets along irrigation ditches and reservoirs, in gardens, graveyards, 
ruins, etc. In Bairam-Ali, on the lower Murgab, jackals colonized and 
dug holes in planted groves of white acacia in a very populous place 
(V.G. Heptner). In Middle Asia, the jackal is, to a considerable degree, 



145 



Fig. 41. Biotope of jackal — ^tugai on the Murgab at Sultan-bent. Turkmeniya, 

10 June 1962. Here also the spotted [steppe] cat and wild boar are found. 

Photograph by A. A. Sludskii. 

a synanthropic animal. It is especially abundant in very old oases, such 
as Tedzhen, Pendin or Merv on the Murgab, Bukhara on the Zeravshan, 
and Khorezma on the Amu-Dar'ya. In connection with irrigation and 
utilization of deserts and the establishment of new oases in the 30's— 
50's of the present century, rapid colonization of jackals was observed 
in the new regions lying in the Vakhsh— Pyandzh interfluve, in the 
Golodnaya steppe of Uzbekistan and in other places. A further increase 
in the number and dispersion of jackal may be expected. 

The jackal avoids waterless deserts, being encountered there only 
at their very edge. As a rule, it does not live in the Karakum and 
Kyzylkum, but is met with in significant quantities along the borders 
of the sandy deserts, adjacent to the Murgab and Amu-Dar'ya oases 
(Salikhbaev, 1950; Stal'makova, 1955), where it comes out from the 
tugais and thickets in order to hunt gerbils. In Repetek, situated only 
70 km away from the Amu-Dar'ya, jackals were not observed 
(Stal'makov, 1955). However, in the fall of 1957 and 1958, they 
appeared here (Sapozhenkov, 1960). They appear in the Asmantai sands 



146 



in the Ustyurt, migrating from the delta of the Amu-Dar'ya and from 
the shore of the Aral Sea. In foothills and mountains they are confined 
to tugai shrubs along ravines, and in bushes along rivers and gullies. 
In the Himalayas, it ascends somewhat higher than 1000 m above sea 
level. 

The presence of jackals in one or another region and their choice 
of habitats are determined by the abundance and availability of prey, 
the presence of water and dense shrubs which serve as shelters both for 
the jackals themselves and for the animals which they hunt. It is espe- 
cially abundant there where water bodies incur no prolonged freezing 
period and where it is possible for waterfowl to overwinter. 

Jackal can withstand low temperatures down to -25° [C] and even 
-35°, but it is not adapted to live in very snowy lands. In the Trans- 
Caucasus, in severe winter with much snow (1949— 1950), jackals could 
travel only on roads and paths made by humans and large animals 
(Vereshchagin, 1951). 








Fig. 42. Habitat of jackal on tlie iviuigal). lugai oi turanga and tamarisl< at 
Sultan-bent. Turkmeniya. 10 June 1962. Photograph by A. A. Sludskii. 



147 

Food. The jackal is a predator-scavenger but, at the same time, it 
readily eats all available carrion and refuse, and feeds on large quan- 
tities of plant food, being, in some seasons, omnivorous. 

In the Caucasus and Trans-Caucasus, it mainly hunts hares and 
mouse-like rodents, as well as pheasants, francolines [F. francolinus] 
(when they were numerous), ducks, coots [Fulica atra], moorhens 
{Gallimila chloropus] and passerines. Birds suffer greatly from it at 
the time of autumn migration and when wintering. It frequently attacks 
domestic birds, including turkeys, more rarely lambs, sheep and goats 
and one even attacked a newborn domestic buffalo calf (Dinnik, 1914; 
Satunin, 1915). They willingly eat lizards, snakes, frogs, fish, molluscs, 
insects, floating dead fish and various [kinds of] carrion. It frequently 
feeds on fruits such as pears, hawthorn, dogwood, and "cones" of 
Mespilus germanicus. It chooses sweet sorts and avoids sour. In gar- 
dens, it destroys many grapes, eating watermelons, muskmelons and 
nuts (Dzhanashvili, 1947). In 26 feces samples collected in summer, 
1950, in the Ismaillinsk region (Azerbaidzhan), mammals were found 
in 90.0% of the total sample, birds — 27.0%, reptiles — 23.1%), 
1 1 3 amphibia — 1 5.4%, insects — 19.2%, plants — ^26.2% and carrion — 1.1% 
(Rukovskii, 1953). In 12 feces samples from Lake Shil'yan 
(Azerbaidzhan) mouse-like rodents were found in 3.3% of the samples, 
water vole [Arvicola] — ^25.0%, European hare [Lepus europaeus] — 
25.0%, carrion (wild boar, dog) — 16.7%, coot — 8.3%, discarded 
fish — 8.3%, insects (locust) — 40.1% and squash seeds — 16.7% (Pavlov, 
1953). 

In severe winters, when water bodies freeze, jackals cause great 
losses among nutria, coots and ducks. In the winter of 1948/49 in 
Trans-Caucasus, nutria occupied first place in the jackal diet (Table 4). 

In such winters, mouse-like rodents, small birds and other foods 
play a small role in the jackal ration. In winter, in the presence of 
abundant prey the jackal kills more animals than it can eat, and caches 
excess food. Jackals feed in a similar manner in Middle Asia and 
Kazakhstan (Table 5). Rodents occupy the principal place and then 
birds (20.0%). 

Some role is also played by fish and arthropods, of which locusts 
and darkling beetles are more frequently eaten (Chemyshov, 1954). In 
some instances the stomachs of jackals are entirely filled with locusts 
(Flerov, 1935). 

In the nutritional ration of the jackal, wild and cultivated plants are 
of serious significance. Thus, the stomach contents of 23 jackals caught 



148 



Table 4. Winter foods of jackal in Trans-Caucasus in the severe winter of 
1948/49 (in % of the total number of samples investigated) (Pavlov, 1953) 



Food designation 


Gagrsk region 


Kurdamirsk 


region 




Abkhazsk ASSR 


Azerbaidzhan SSR 




Before freezing 


During freezir 
104 samples 


ig period 




30 samples 


17 samples 


Nutria 


26.6 


89.4 


47.0 


Water vole 


3.3 


0.9 


11.6 


Brown rat 


13.3 


2.8 


5.9 


[Rattus norvegicus] 








Voles 


— 


— 


5.9 


Asia Minor gerbil 


— 


— 


5.9 


[Meriones tristrami] 








Small mouse-like rodents 


26.6 


18.2 


— 


Birds 


36.6 


76.9 


64.7 


Including: 








coot 


— 


— 


29.3 


mallard 


— 


— 


23.5 


grebe 


— 


— 


5.9 


passerines 


— 


— 


5.9 


domestic fowl 


10.0 


— 


— 


Fish 


6.6 


— 


— 


Insects (beetles) 


— 


1.9 


5.9 


Carrion of domestic animals 20.0 


— 


— 



in October-January in tugais on the Vakhsh consisted completely, or 
in large part, of the fruit stones of wild stony olive {Elaeagnus 
angustifolia) (Chemyshev, 1948). In spring, jackals dig out and eat 
plant bulbs (Flerov, 1935; Chemyshev, 1954), and later, roots of wild 
sugar cane (Table 6). 

Jackals living near the edge of the Karakum venture out into sandy 
areas to hunt, where they catch gerbils, lizards and snakes. In the 
period of drying of canals and irrigation ditches, they feed on fish and 
frogs, and hunt muskrats. In tugais and gardens, jackals eat fruits of 
wild olive (dzhida)*, mulberry and dried apricots, as well as water- 
melons, muskmelons, tomatoes and grapes, to which they cause 
appreciable harm. 

In the lower Amu-Dar'ya, in the stomachs and feces of jackals, 
were found remains of tolai hare, complex-toothed rat [Nesokia indica]. 



*Dzhida is a local name for wild (stony) olive — Sci. Ed. 



149 



Table 5. List of jackal foods in southwestern Tadzhikistan based on analysis 

of gastro-intestinal tract samples (60 specimens) and feces (175 specimens) 

(Chernyshev, 1958) 



Type of food 


% of 235 


Type of food 


% of 235 




samples 




samples 


Animal food 


78.8 


Reptiles and amphibians 


2.8 


Mammals 


36.8 


Reptiles unidentified 


0.8 


Mammals, unidentified 


2.0 


Lake frog 


2.0 


Carnivores 


0.8 


Fish 


6.8 


Badger 


0.4 


Fish, unidentified 


2.0 


Marbled polecat 


0.4 


Gambusia 


3.6 


[Vormela peregusna] 




Common саф 


1.2 


Rodents 


31.2 


Arthropods 


14.4 


Rodents, unidentified 


6.0 


Insects (majority are 




Tolai hare [Lepus tolai] 


11.2 


beetles, and others) 


9.6 


Field rat* 


3.2 


Plant food 


32.6 


Turkestan rat 


1.6 


Melon culture (musk- 




[Rattiis turkestanicus] 




melon, watermelon) 


6.4 


Red-tailed gerbil 


2.4 


Wild olive fruits 


14.4 


[Meriones erythroiinis] 




Wild cane sugar 


4.0 


House mouse [Mus musciilus] 


6.4 


(root, stems) 




Mole-vole [Ellobius sp.] 


0.4 


Bulbiverous plants 


4.4 


Birds 


20.0 


Nightshade fruit 


1.6 


Birds, unidentified 


2.8 


Wolfberry fruit 


1.2 


Pheasant 


10.8 


Wheat 


0.8 


Duck 


2.4 


Carrion 


10.4 


Coot 


0.8 






Passerines, unidentified 


3.2 







''Species not clear — Sci. Ed. 



gerbils, pheasant and other birds, insects (locust and beetles) and rarely 
plants (Salikhbaev, 1950). Where there are muskrats and nutrias, the 
jackal destroys a great number of these rodents (Pokrovskii, 1953; 
Kostin, 1956). 
1 1 5 They visit the sea shore where they pick up the dead fish, wounded 

waterfowl, molluscs, and feed on parts of marine animals discarded by 
processors. They also feed near slaughterhouses, dumps, and cattle 
burial places. In southern Dagestan in the 20's, jackals regularly fed 
near the railway lines, picking up food remains thrown out from pass- 
ing trains by passengers. They were so accustomed to finding edibles, 
wrapped in paper, that hunters placed poisonous baits along the railway 
lines, wrapping it in newspaper (Heptner and Formozov, 1941). 



150 

14 Table 6. Seasonal variation in the list of principal foods of the jackal in 
southwestern Tadzhikistan (Chernyshev, 1958) 



Name of food 




Season of year 






Winter 


Spring 


Summer 


Autumn 




XII, I, И 


III-V 


VI-VIII 


IX-XI* 


Rodents 


3' 


4 


3 


3 


Birds 


3 


4 


4 


3 


Reptiles and amphibians 


I 


2 


3 


2 


Fish 


1 


2 


3 


3 


Arthropods 










(majority are insects) 


1 


3 


4 


3 


Wild plants 


3 


3 


3 


4 


Cultivated plants 


3 


3 


3 


4 


Carrion, refuse 


3 


3 


3 


3 



*Designates months of the year — Sci. Ed. 

'Frequency of occurrence of food: very frequent — 4; frequent — 3; rare — 1\ 
very rare — 1 . 

115 In summer the jackal, apparently, needs to drink regularly, since at 

that time they always stay near water and are not found in waterless 
deserts. In southeastern Turkmeniya, when considerable sections of the 
Kushka [river] dry up, jackals, together with wolves and foxes, dig 
holes in the bottom of the dry channel and drink the ground water 
collected in them (Yu.K. Gorelov). Eating of watermelons, muskmelons 
and grapes is also linked with the slaking of thirst. 

Home range. In southwest Tadzhikistan, daily movements of jack- 
als sometimes comprise 8—12 km, i.e. the home range of one family in 
tugai has a radius of up to 12 km. In Tigrovaya Balka Preserve, in an 
area of 6000 hectares, from 10 to 23 jackal families were counted in 
different years, i.e., one family occupies an average of from 260 to 600 
hectares (Chernyshev, 1954). In Georgia, at Lake Inkit, in the severe 
winter of 1948/49, seven families of this carnivore occupied an area of 
3x3 km (M. Pavlov, 1953). The average area for one family there was 
130 hectares. 

In case of food shortages in some winters, and also after large fires 
in tugais and reeds, jackals in search of prey were obliged to undertake 
movements of 40—50 km, regularly appearing in cultivated lands and 
villages.** 



4n Russian original, kishlak. a Turkic word for village — Sci. Ed. 



151 

Burrows and shelters. In the Caucasus and Trans-Caucasus, the 
female usually gives birth in a burrow it has dug together with the male, 
or she occupies the abandoned dwellings of fox and badger. The burrow 
is dug some days before delivery, and the female and male take turns 
digging. It is located in thick shrubs, on the slope of a ledge or gulley, 
or on a flat surface. The burrow is simple in structure, since it possesses 
one opening which runs to a depth at a small angle. The length of the 
burrow is about 2 m, while the nest chamber is located at a depth of 1 .0- 
1 .4 m. In Dagestan and Azerbaidzhan, jackal litters also were repeatedly 
found in the hollows of large fallen trees (Satunin, 1915; Dzhanashvili, 
1947). In Azerbaidzhan, lairs with pups were found under the roots of 
trees and under large stones on river banks (Dzhanashvili, 1947). 

The litter remains in the burrow for 2—2.5 months after which the 
female leaves the lair with the young and begins to lead a roving way 
of life. Regardless of the time of reproduction, the lair is built in dense 
thickets of spiny bushes (blackberry, Jerusalem, or Christ thorn) or 
reeds, and rarely in caves, crevices in rocks, hollows, old burrows, 
water channels and other secluded places. 

In Middle Asia, jackals construct lairs in the dense thickets of 
tugais, and burrows are not dug (Chemyshev, 1954), but they have 
been described for the Vakhsh tugais (Flerov, 1935). These burrows 
usually have short, straight passages, dug under the roots of turanga or 
wild olive, or directly in dense thickets of tamarisk and Nitraria sp. 
One of these burrows, with a length of three meters or more, went 
obliquely downwards for some distance and widened into the lair itself. 
In front of the entrance there was a quite large heap of earth which was 
greatly trampled down in the middle. At a distance of 140-200 m two 
other holes, similar to the first one, were found. From all the holes, 
well-packed paths extended through the tugai, which led to a common 
network of trails. Surrounding the holes were many feces, partially 
covered with earth. 

In the tugais and cultivated lands of Tadzhikistan, jackals construct 

lairs in the turf of a gigantic grass — plume grass [Erianthus], less 

16 frequently in Nitraria sp., shrubs and in openings in reeds. There, 

where vegetation may have been destroyed, jackals live in holes. The 

female, male and young live in one lair (Chemyshev, 1954). 

Daily activity and behavior. The jackal basically follows a cre- 
puscular and nocturnal way of life, leaving its diurnal shelter before 
sunset and returns with the light or even later in the morning. They also 
often hunt by day. One may judge whether it is leaving to hunt by the 



152 

character of its howl. When one animal begins to howl, the others soon 
answer. Howling is, apparently, also connected in part with the weather. 
Cases have been noted, when, in case of change in weather, jackals 
break out in a prolonged and continuous howl. Jackals accompanied 
with howls the ringing of church bells; the young start to howl when 
they hear the sound of a siren, or whistle of steamship or train engine. 
They usually howl in the evening and at dawn, but sometimes during 
the day, even at midday. 

The jackal is a daring and brave animal, and in places where it is 
not pursued, less cautious and not afraid of people. It is easily caught 
in traps or envenomed with poisoned bait. Many instances are known, 
when, within sight of people, it attacked a domestic fowl, and regard- 
less of the shouts and pursuit, takes it off. From field camps and from 
houses it drags off various objects, which it thinks to be edible, such 
as leather, boots, rags, venturing to steal them just a few steps from 
sleeping people. 

Jackals usually hunt singly, and less frequently in pairs and in [a 
group of] 3-4 and only rarely do they form small packs. In the Trans- 
Caucasus, at the end of summer and in autumn, packs of 8—12 jackals 
were observed including, apparently, two or more families (Dzhanashvili, 
1947). When embarking on a hunt, the jackal jogs along at a trot and 
frequently stops to sniff and listen. On spotting prey, it stealthily con- 
ceals itself, and quickly approaching, pounces and quickly makes a kill. 
When hunting in pairs or packs, they run in parallel, overtaking the 
prey together. For example, when hunting nutria or muskrats, they run 
along both banks of narrow streams or channels, driving the frightened 
prey from one to another. When lying in wait for its prey, it conceals 
itself near a path or waterhole. In times of drought, they concentrate 
near the drying lakes and channels and catch animals and birds on the 
shore or in shallow water; they also catch fish there. During freeze-up, 
they hunt waterfowl, nutria and muskrats coming out on the edge of the 
ice or bank. They attack sleeping flocks of birds (ducks, little bustards 
[T. tetrax]) and especially many of them are destroyed in severe win- 
ters when they are exhausted from starvation. In autumn, when the 
canals and irrigation ditches are prepared for cleaning and are dry, 
jackals together with foxes, jungle cats, badgers, crows, and magpies 
successftilly hunt fish and frogs that have collected in the remaining 
pools of water, as well as muskrats forced to walk over significant 
section of dry land. When water bodies freeze, in trying to catch 
muskrats, they destroy their houses. 



153 

The jackals live in pairs, but are also found either singly, or in 
pairs and families of 4-5 individuals. 

Jackals, when taken young, quickly become tamed, following their 
owner everywhere, and running to his call. They behave as an affec- 
tionate dog would. Even adult animals can be tamed. In some places 
the dogs do not disturb jackals and both animals could be observed 
together on one garbage heap. 

Seasonal migrations and transgressions. Regular seasonal migra- 
tions are not characteristic of jackals. They usually wander only in 
various directions searching for places with food in which they concen- 
trate. Thus, in autumn they gather on the shore, where they pick up bird 
remains. On sea coasts they are attracted by the carcasses of dolphins 
or seals thrown out by processors. During fall, the number of jackals 
in the tugai grows due to the arrival of animals from neighboring 
cultivated lands. In winter, they wander into human settlements or 
intrude deep into the desert. During such wanderings, they cross 50— 
100 kilometers or even more. 

While not truly regular migration, at the same time jackals often 
settle at great distances from their usual range, and appear in regions 
in which they were only a long time ago, or where they are completely 
unknown (see above). In 1920-1922, when many cattle died of starva- 
tion, jackals appeared in considerable quantity in western Kazakhstan, 
especially in the valley of Ural river; in the north they reached nearly 
to Orenburg (Sludskii, 1939). The jackal may appear around Orenburg 
from the delta of the Amu-Dar'ya, passing along the shore of the Aral 
Sea, through Ustyurt to the Emba and Ural; this way constitutes, in a 
straight line, about 1000 km. 

In the 30's of the present century, the jackal was rare in the middle 
course of Syr-Dar'ya (Aris' region), but in 1942—1945, it became 
common there, reached eastward to Chimkent, appeared in the Boroldsisk 
mountains in the upper Bugun 150 km north of Chimkent (Sludskii, 
1953). It remained in the latter region until the 50's. 

In the winter of 1948—1949, in Kustanaisk district, after an ex- 
tended continuous snow cover and massive death of cattle on winter 
pastures, jackals appeared in the summer of 1949, in Dzhangil'dinsk 
and Amangel'dinsk regions of this district, on the Turgai river and at 
Lake Sarykopa, and became common. Through 1952, their skins were 
received in trade. The appearance of jackals in 1949 in the Turgais 
steppes was accompanied by a massive inflight to this area of brown 
[Gyps fulvus] and black [Aegypius monachus] vultures which were 



154 

also attracted by the abundance of carrion. On the Turgai river, jackals 
could pass from the delta of the Amu-Dar'ya along the western shore 
of the Aral Sea and the Bolshoi Barsuki sands, or also from the Syr- 
Dar'ya, below Aris', across the lower course of this river and the 
Priaral'sk Karakum sands. In the first event they might have gone 
about 1050 km, and in the second — ^about 1125 km. 

In the severe winter of 1950/51 in southern Kazakhstan, jackals 
appeared at the northwestern edge of the Betpak-Dala desert, in the 
Dzhetykonur sands, having penetrated there from the Syr-Dar'ya up- 
wards along the Sary-su river, and they remained there in limited 
numbers until 1954; several animals, in that winter, succeeded in reach- 
ing even the upper Sharubai-Nury river 40—50 km southwest of 
Karaganda. The route thence from the Syr-Dar'ya is about 800 km. In 
the lower Chu river, the jackal was encountered at the end of the past 
century, then disappeared and again appeared in 1917. Later, individual 
animals were caught there in 1927, 1936 and 1948. 

In October 1924, a female of a pair was caught 60 km from 
Tambov (Ognev, 1931). In order to reach Tambov from the lower 
Terek along the Caspian Sea coast and upwards along the Volga and 
its tributaries, jackals must have gone about 900 km. The appearance 
of carnivores in new regions usually coincides with the mass death of 
ungulates from starvation. It is not clear how jackals and vultures fmd 
separate regions where there is much carrion. 

In Bulgaria, jackals also sometimes migrate to the north and northwest, 
appearing near Sofia, at the city of Kotel in Asenovgradsk region, at the 
city of Dimitrova, and also intruding into Romania, Yugoslavia and Hun- 
gary (Atanasov, 1953). In the winter of 1952/53 they appeared in Slovenia, 
following, apparently, the herded sheep. Prior to this winter, jackals in 
Yugoslavia were only found in Dalmatia and Macedonia (Berlikh, 1955). 

Reproduction. Jackals live in pairs, united apparently, for life, 
until one of the partners dies. The male helps in digging the burrow and 
raising the young. In southwestern Tadzhikistan, out of 82 jackals, 39 
were males and 43, females (Chemyshev, 1958). 

In the Trans-Caucasus, estrus begins in the first half of February, 
and in warm winters, at the end of January. The mating period conti- 
nues for 26-28 days (Dinnik, 1914; Satunin, 1915; Dzhanashvili, 1947). 
In Tadzhikistan and Uzbekistan, the rut manifested itself at the same 
time also, and extended to the first days of March (Chemyshev, 1954; 
Kashkarov, 1931; Salikhbaev, 1950). In Bulgaria, it extends from the 
middle of January to mid-February (Atanasov, 1953). 



155 

18 Gametes in adult males and females do not mature at the same 
time. Active spermatogenesis in males appears 10—12 days earlier than 
estrus in females. During the mating period, testes weight increases 
three times.* In Tadzhikistan, average weight of testes in January 
was 6.1 g, in February — 13.3, in March — 13.0, in April — 7.7 and in 
August — 4.4. In a male, caught on 6 March 1948, with noticeable 
degradation of the testes, sperms were still detected. Ovaries increase 
insignificantly in the estrus period (from 0.7 g in January to 0.8-0.9 g 
in February— March) (Chemyshev, 1954). Estrus continues 3^ days. If 
the female is not mounted during this period, her receptive condition 
will be interrupted for 6—8 days, after which estrus returns and again 
continues 3-4 days. If the female does not become pregnant by that 
time, estrus will not recommence until the next year. Mating can be 
observed at any time during the diurnal cycle and is accompanied by 
an attachment [mating tie] which lasts for 20-45 minutes (Dzhanashvili, 
1947). 

Jackals nowl particularly often during the mating period. The fe- 
male coming into heat for the first time is pursued by several males, 
and quarrels develop among them. The duration of pregnancy is 60-63 
days (Bogdanov, 1873, Pocock, 1941, Brink, 1957); according to 
observations at Tbilisi zoo — 62-63 days (Dzhanashvili, 1947). In 
Tadzhikistan a considerable number of barren females (5 out of 15) 
was noticed in different years (1948, 1950). In all of these barren 
females, helminths were found in great quantities (Chemyshev, 1954). 

In the Trans-Caucasus, birth of the young usually takes place from 
the end of March to the last days of April. The earliest parturition was 
noted on 28 March 1937 and the latest, on 25 April 1931 (Dzhanashvili, 
1947), but sometimes females deliver in the first half of May (Satunin, 
1915). In Tadzhikistan, Uzbekistan and Bulgaria parturition occurs in 
April and the beginning of May. Only once, a female gave birth to 8 
pups on June 1 1th (Chemyshev, 1958; Atanasov, 1953). Only one litter 
per year is produced. 

In the Trans-Caucasus, the number of young in the litter ranges 
from 3 to 8 (Dinnik, 1914; Satunin, 1915; Markov, 1934; Dzhanashvili, 
1947). Satunin (1915) dug out several burrows, in one finding 5 pups, 
and in the remainder, 4. According to observations in nature and in 
zoos, the litter usually contains 5 pups, rarely 3^ or 6—8. In 
Tadzhikistan, there are 3—7 (average 4.8 pups; Chemyshev, 1958). In 

*The weights given in the next sentence suggest that this should read "two times" — 
Sci. Ed. 



156 

Uzbekistan, there are 2-8 pups in a litter (Kashkarov, 1931; Salikhbaev, 
1939, 1950). In the majority of regions in Bulgaria, there are 4-7 pups, 
but in Michurinsk region, only 3—5 (Atanasov, 1953). In India, a litter 
contains an average of 4 pups (Blanford, 1888). 

The female nurses the young for 50—70 days (Caucasus; 
Dzhanashvili, 1947), in Tadzhikistan up to 90 days. The lactation 
period in some females ends in the middle of July and in others, in the 
first days of August (Chemyshev, 1958). The young begin to eat meat 
starting at the age of 15—20 days. Only in rare instances does the 
mother feed the pups regurgitated food. At the end of the lactation 
period, the female drives the young away from her by growling and 
biting. The latest-bom remain with the female until the first half of 
autumn, after which they leave her and start leading an independent 
life, either singly or in groups of 2-A individuals. 

Sexual maturity, based on observations in the Tbilisi zoo, takes 
place in females at an age of 1 0— 1 1 months, but in males at the age 
of 21— 22 months. In males lO-l 1 months old, the testes are undevel- 
oped and contain no sperm (Dzhanashvili, 1947). In Tadzhikistan, males 
are said to become sexually mature at the age of 10—11 months 
(Chemyshev), a statement which is not true. Duration of life is 12—14 
years (Brink, 1957). 

Growth, development, and molt. Pups are bom with a pelage of 
soft fiir, the color of which varies from light-gray to dark brown. At the 
age of one month, the juvenile pelage is replaced. At first, guard hairs 
grow on the head, back and mmp, then on the anterior and posterior 
19 limbs and, finally, on the tail. Fur color of the young changes, becom- 
ing reddish with black speckles. Young caught by the end of August 
and the beginning of September hardly differ from adults in fur color. 
By October and November, the young already have full winter fur 
(Chemyshev, 1958). 

Young are bom with closed eyelids. In the Tbilisi zoo, in 39 pups 
(78%) the eyelids opened on the 9th and in the remainder — on the lOth— 
1 7th day. Eye-opening is delayed in weak and sick pups (Dzhanashvili, 
1947). In Tadzhikistan, based on observations of 9 pups, eyelids opened 
on the 8th— 1 Ith day (Chemyshev, 1958). Ears in healthy pups open on 
the 1 0th— 13th day and become erected only on the 25th— 30th day. 

The last [3rd] and middle [2nd] deciduous incisors in the upper jaw 
first erupt on the 1 lth-14th day. The inner [1st] deciduous incisors of the 
lower jaw are the last to empt, on the 17th-l9th day. At that time, the 
deciduous canines appear. The replacement of the deciduous by permanent 



157 

incisors is completed at about the age of 4 months, and the third and final 
molar of the lower jaw erupts at an age of about 5 months. The times of 
eruption of deciduous teeth and the order of their replacement by the 
permanent ones is related to lactation, protracted feeding of the pups, and 
the time of their transition to an mdependent life. 

Young jackals grow quickly. At the age of two days their weight 
is 201-214 g, at one month, from 560-726 g, and at 4 months, 2700- 
3250 g. Young caught at the beginning of September weighed 4200-4370 
g, and by the beginning of October, their weight reaches 3/4 of adult 
weight (Chemyshev, 1958). 

The jackal molts twice yearly — ^in spring and in autumn. In the 
Trans-Caucasus and Tadzhikistan, the spring molt begins in the middle 
to end of February, in warm winters, in the middle of that month; in 
cold winters, it starts in mid-March and ends at the beginning to middle 
of May. In healthy animals, pelage replacement extends for 60-65 days 
in spring. Weak and sick animals do not succeed in completing molt 
during the entire summer; at the beginning of winter, half of the coat 
consists of spring— summer hair and half is winter fur (Dzhanashvili, 
1947). 

Spring molt starts with the head, the anterior and posterior limbs, 
then spreads to the sides, back, chest, belly and rump, and the last to 
molt is the tail. After spring molt, only scattered coarse guard hairs 
remain on the skin and underfiir is absent. Autumn molt occurs from 
the middle of September. Shedding of summer pelage and growth of 
winter guard hairs and underfiir take place simultaneously. Autumn 
molt starts with the rump and tail and spreads to the back, sides, belly, 
chest, limbs and head. Full winter fiir has grown out by the end of 
November (Chemyshev, 1958). In the Caucasus, the skin is prime from 
November to March (Vereshchagin, 1947). 

Enemies, diseases, parasites, mortality, and competitors. Earlier, 
enemies of the jackal everywhere were the tiger and leopard. At the 
present time, the first is extinct in the jackal's range, and the second 
is extremely rare. The leopard hunted jackals especially frequently. At 
the same time, jackals regularly fed on the remains of these cat's prey, 
and once, in Tigrovaya Balka, they ate a dead tiger. The wolf also 
should apparently be considered an enemy. 

Its serious competitors everywhere are the jungle cat, red fox, wolf, 
and in the Trans-Caucasus, the forest wildcat [Felis sylvestris], and to 
a lesser degree, the recently acclimatized raccoon [Procyon lotor] as 
well. In Middle Asia and Kazakhstan, the steppe wild cat {Felis libyca] 



158 

is also a competitor. The small number of red foxes occurring in tugais 
may explain the abundance of the jackal there (Chemyshev, 1958). At 
the same time, in deserts and foothills, red fox and wolf apparently oust 
the jackal. Usually, where there are many wolves, jackals are absent, 
and vice versa. 

Among the infectious diseases of jackal, carnivore distemper and 
rabies are established (Sludskii, 1954; Chemyshev, 1958). Epizootics 
120 of carnivore distemper in the winter of 1948-49 in Tadzhikistan 
(Tigrovaya Balka) affected not only jackals, but was also observed 
among shepherd and hunting dogs, the majority of which died. In south- 
western Tadzhikistan, of 9 jackals examined, leishmania was found in 
liver and spleen smears of three, and was supposedly identified as 
Donovan's leishmania, which induces visceral leishmaniasis in humans. 
Jackals infected with leishmania suffered little from this infection 
(Latyshev and others, 1947). 

Jackals are heavily infested with helminths. In southwestern 
Tadzhikistan, 15* species of cestodes, roundworms and acanthocepha- 
lans were discovered: Sparganum mansoni, larval form of 
Diphyllobothrium mansoni, Taenia hydatigena. Taenia pisiformis. 
Taenia ovis, Hydatigera taeniaeformis, Diphylidium caninum, 
Mesocestoides lineatus, Ancylostoma caninum, Uncinaria 
stenocephala, Dioctophyme renale, Toxocara canis, Toxascaris leonina, 
Dracunculus medinensis, Filariata gen. sp., [and] Macracanthorhynchus 
catulinum. Jackals infect water bodies with the parasitic worm larvae 
{D. medinensis) and may be responsible for dracunculosis in humans 
(Chemyshev, 1954). Moreover, they play some role in the spread of 
coenurosis of sheep and large homed cattle. In Uzbekistan, 10 species 
of helminths were found in the jackal, of which, in addition to 7 species 
mentioned for animals in Tadzhikistan (No. 6-12, see above), were 
found: Spirocerca lupi, Rictularia affinis and R. cahirensis (Irgashev, 
1958). 

Jackals in Tadzhikistan are infested with ticks in April-September, 
while in November— Febmary, these parasites are only found in few 
individuals and in small numbers. The following species of ticks were 
found: Ixodes sp., Rhipicephalus turanicus, R. leporis, R. rossicus, R. 
sanguineus, R. pumilio, R. schulzei, Rhipicephalus sp., Hyalomma 
anatolicum, H. scupense, H. asiaticum, Hyalomma sp. Most frequently, 
R. sanguineus parasitizes the jackal in great quantities. The role of 
jackals in nourishing and spreading ticks is great. Four flea species are 

*16 are listed — Sci. Ed. 



159 











Fig. 43. Track of jackal on silt. "Tigrovaya Balka", preserve at Pyandzh, 
Tadzhikistan. 15 October 1960. Photograph by A. A. Sludskii. 

known on jackals: Pulex irritans, Xenopsylla nesokiae (a flea of the 
bandicoot rat), Ctenocephalides canis, C. felis and one species of 
louse — Trichodectes canis (Chemyshev, 1954). 

In severe deep-snow winters, jackals often die from emaciation and 
exposure. In Azerbaidzhan, in severe winters, after the large lakes are 
covered with ice, jackals penetrate far into the dense growths of reeds 
and cattails and stay in them during the entire frozen period, not going 
to shore. At times of sudden thaws, they are stranded on the ice and 
die. Thus, in December 1946, when a thaw commenced, the animals 
which were living in the reeds were isolated from the land by water and 
drowned (Vereshchagin and Dyunin, 1949). In the severe winter of 
1948^9 at the Shil'yansk lakes, as a result of sudden thawing, the 
animals became isolated from the shore and drowned at the borders of 
the impassable flooded reeds in their attempts to reach dry land. Other 
cases were observed when jackals, failing to reach the shore, remained 
starving for a long time on intact sections of ice (Pavlov, 1953). Near 
the Caspian Sea jackals sometimes die in great numbers in autumn and 
winter moryan* which quickly fill coastal thickets, and in times of 
floods. Possibly, they also suffer from fires in the turgais and reeds. 

*Violent on-shore storm — Sci. Ed. 



160 

Population dynamics. These are little studied. Judging by the data 
of fur buyers, the number of jackals is subject to considerable fluctua- 
tion. Thus, in Turkmeniya, fur purchases varied by nearly 3-4 times 
during the period from 1950 to 1956 (1400-5500 items). They fell 
especially greatly in 1954 — to 1430 — ^following the severe winter of 
1953-54. Fur purchases considerably increased in 1956, reaching 5427 
skins, following 1955, a year which was favorable for reproduction 
because of abundant food. 

In Tadzhikistan, minimal fur purchases during the ten years was in 
1951 — ^258 pieces after the severe winter of 1950-51, and the maxi- 
121 mum, as in Turkmeniya in 1956 — 1403 skins. In Tadzhikistan, fur 
purchase, and consequently the number of jackals, fluctuated approxi- 
mately by six times. In the lower Amu-Dar'ya in the decades of the 
30's-50's the catch of jackal skins varied by 2 times and was also 
related to the degree of severity of winter. In the south of Kazakhstan, 
jackals were relatively common in the 20 's and 40 's of the present 
century, during periods when there was much carrion in the desert. 

In 1926-1929 in Kazakhstan, from 100 to 250 jackal skins were 
obtained; at the beginning of the 30's, this number dropped to 25-30 
pieces; after which a few individuals were caught; but in 1942—1945 
the number of jackals started to increase. By 1947, jackals were com- 
mon in the most southerly regions. In this year, 104 jackal skins were 
tanned in Kazakhstan. In the following years, their number again 
dropped. However, these changes were mainly connected with their 
migrations from the south. 

Field characteristics. The jackal is half the size of the desert wolf 
and nearly twice as large as the desert red fox, and is high in the limbs. 
Ears are erect, but more blunt than those of foxes. Tail short and 
1/3 the body length. 

Feces are sausage-like, and are found in noticeable piles, as the animal 
leaves them in one and the same place for several times. It often howls. The 
howling is similar to a wail, and can be rendered by the sounds "Ai-yai! 
Ai-yai!" — high, whimpering. It is similar to that of dogs, but is higher and 
more varied and is differentiated by a "plaintive" intonation. To the howl 
of one jackal, those nearby answer (A.S.). 

Practical Significance 

The jackal is a fur-bearer, but its fur is of low value, characterized by 
sparse, coarse pelage and has a monotonous unbeautifiil color. Already 



161 

in the last century the jackal was utilized (Silant'ef, 1898). In the 
Trans-Caucasus, in the Zakatal area, 300 jackals were captured in 
1896, and in the B[olshoi] Trans-Caspian district, the jackal was hunted 
in many jurisdictions, and in Mervsk, in the eighties, about 200 animals 
were caught yearly. The total taken in Russia in that period was more 
than ten thousand jackals. Jackal skins were sent to the Nizhegorod fair 
(they were not sent to other fairs) in the following numbers: 

Year 1880 1881 1883 1884 1885 1886 1887 

Number of skins (in units) 8000 10,000 3000 4000 4000 3000 2800 

Year 1888 1889 1890 1891 1892 1893 

Number of skins (in units) 3000 3000 3500 4000 4000 3000 

In the decade of the 20's in this century, the following numbers of 
skins of this animal were tanned in Trans-Caucasus (Markov, 1934): 

122 Years 1924- 1925- 1926- 1927- 1928- 1929- 1931 1932 

1925 1926 1927 1928 1929 1930 

Number of skins 4386 6159 4027 11793 9334 13635 11819 13384 

In Middle Asia and Kazakhstan lesser amounts of skins were pre- 
pared (Tables 7, 8). 

In the entire Soviet Union, at the beginning of the 30' s of the 
current century, about 20—25 thousand were tanned yearly (Naumov 
and Lavrov, 1941; Kaplin, 1960). However, the stocks of this animal 
were significantly underutilized. In 1940, 10.4 thousand fiirs were pro- 
cessed. At present, it is possible to increase the size of the jackal take 
by a minimum of three-fold. The world production of jackal skins in 
1907-1910 was 20-25 thousand items (Asia 5.0, Europe 5.0, Africa 
10.0; Kaplin, 1960). Previously, the majority of skins tanned in the 
USSR were sent abroad, mainly to the USA. Since 1949, the skins have 
been utilized inside this country. 

The skins are not graded according to a fur standard, though those 
obtained from the Transcaucasus possess coarser fur of reddish-brown 
color, while those from Middle Asia differ in their softer paler fur. 
They are divided into four sorts, and used in the manufacture of cheap 
collars, women's coats and dokhas [fur-coats]. They are usually dyed. 
Suede is prepared from the skins of grade IV. 

A special harvest of jackals does not exist. The animal is caught 
incidentally to the hunting of other animals, by means of traps or by 



162 



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163 

shooting while on drives. When traps are employed, they are set near 
carrion, on garbage dumps (in such places the beast pays little attention 
to the odor of metal) and in shallows along shores; as a bait, a large 
live fish is tethered on a kiikan* (string). 

In the Trans-Caucasus, jackals are hunted with large fishing hooks 
which are suspended from a metallic cable or wire at a height of 75— 
100 cm from the ground, in such a manner that the predator can reach 
it only by jumping. The hook is baited with a piece of carrion. On 
seizing the bait, the animal is hooked by the lip or jaw (Markov, 1934). 
The jackal is snared by wire nooses placed on their uail. Poisons are 
also utilized (see above concerning food). 
123 The jackal is a serious pest of the agricultural and hunting econo- 

mies. In the Trans-Caucasus, "jackals must be considered the most 
harmful destroyers of game" (Dinnik, 1914), causing the most severe 
losses to the hunting economy, not excepting the wolf (Markov, 1934). 
They destroy, in large quantities, newborn roe deer, hares, and nutria 
as well as pheasants, francolins, gray partridges, little bustards and 
great bustards during overwintering, and waterfowl (Satunin, 1915; M. 
Pavlov, 1953; Katubadze, 1956; Burdzhanadze, 1959). The jackal is a 
dangerous enemy of the nutria, completely destroying them in shallow 
water bodies (M. Pavlov, 1953). They are very harmful to the muskrat 
trade. In the Amu-Dar'ya delta, during the winter of 1948-49, in the 
feces of this predator the remains of muskrat constituted 12.3% of all 
examined samples. In particular places, jackals destroyed up to 71% of 
the muskrat houses, of which about 16% froze and became unsuitable 
for occupation (Pokrovskii, 1953). It causes also great harm to the 
muskrat trade in the Khorezma district where muskrat houses are easily 
accessible to it. The jackal also damages the muskrat harvest by eating 
muskrats caught in traps, and skins hung up for drying (Kostin, 1956). 

The jackal is very harmful to poultry and carries off lambs and kids 
(Satunin, 1915). It eats ripe grapes in such quantities that earlier, in the 
Bol. Lenkoran vineyards, owners removed the flowering branches hang- 
ing near the ground because the fruits will in any case be eaten by 
jackals (Satunin, 1915). The jackal destroys watermelon and melon. In 
one melon field in Tadzhikistan, the jackals destroyed, ate and spoiled 
70 ripe melons in one night (Chemyshev, 1958). In the Trans-Caucasus 
and Middle Asia, the jackal, as much as the red fox and wolf, serves 
as a carrier of rabies in nature. 

*Local word? — Sci. Ed. 



164 

Cases are known in which rabid jackals have run into villages, 
where they have bitten man and domestic animals (Bikhner, 1905). The 
value of jackals as eaters of small rodents and insects is inconsiderable. 

The jackal may be killed the year round, until such time as their 
numbers proceed down to a minimum. In all republics, there exist 
decrees permitting their hunting throughout the entire year. (A.S.) 

WOLF 

Canis lupus Linnaeus, 1758 

1758. Canis lupus. Linnaeus. Syst. Nat., Ed. X, 1, p. 39. Upsala, 

Sweden. 
1792. Canis lupus albus. Kerr., Animal Kingdom, p. 137. Lower 

Yenisei. 
1804. Canis lupus communis. Dwigubski. Prodromus faunae Ross., p. 10. 

"Russia" — ^apparently, the northem forested section of European 

Russia (the steppe wolf Dwigubski inserts separately; see below). 
1804. Canis lupus campestris. Dwigubski. Ibidem, p. 10. Cis-Cauca- 

sian steppes. ^ 
1831. Canis pallipes. Sykes. Proc. Zool. Soc. London, p. 101. Deccan, 

India. 
1847. Lupus laniger. Hodgson. Calcutta Joum. Nat. Hist., 7, p. 474. 

Tibet. Nee Canis laniger H. Smith 1840 (domestic dog of Puget 

Sound Indians). 
1863. Canis chanco. Gray. Proc. Zool. Soc. London, p. 94. "Chinese 

Tatary". Tibet." 
1875. Canis anindinaceus. Karelin. Tr. SPb Obshch. Estestvoisp., 6, 

p. 81. South Ural steppe. Nomen nudum. 
1882. Canis lupus var. desertorum. Bogdanov. Ocherki prirody 

Khivinskovo oazisa i pustyni Kyzylkumy, p. 30, Kyzylkum. 
124 1907. Lupus filchneri. Matschie. Wiss. Ergebn. Exped. Filchner nach 

China u. Tibet, 10, 1, p. 153. Si-ning to the east from Kukunor. 
1907. Lupus karanorensis, Matshie. Ibidem, p. 157. Khala-chu in Gobi 

(a little west of Sa-chzhou. V.H.). 
1907. Lupus tschiliensis. Matschie. Ibidem, p. 160. Shanghaiguan' on 

shore of Lyaodunsk gulf 



'For reasons, see later in section "Geographic Variation". 



165 

1911. Lupus altaicus. Noak. Zool. Anz., 35, p. 465. Teletskoe lake 

and Chulyshmansk mountains, Altai. 
1922. Canis lupus var. orientalis. Auct. Dybovski. Archiv Tovar, 

Naukow. Lwowe, 1, p. 350. Forest of Eastern Siberia. Nee 

orientalis Wagner 1841. Nomen nudum. 
1922. Canis lupus var. argunensis. Dybowski. Ibidem, p. 350. Steppes 

of Dauriya. Nomen nudum. 

1922. Canis lupus var. kamtschaticus . Dybowski. Ibidem, p. 350. 
Kamchatka. Nomen nudem. 

1923. Canis lupus cubanensis*. Ognev. Biologicheskie izvestiya, 1, p. 
114. Mountains of Maikop area. Western Caucasus (district of 
present Caucasian preserve, at sources of Kuban tributaries — 
Laba and Belaya). 

1923. Canis lupus turuchanensis . Ognev. Ibidem, p. 115. Turukhan 

territory (right bank of lower Yenisei). 
1926. Canis lupus dybowskii. Domaniewski. Annal. Zool. Musei 

Polonici Hist. Nat., 5, p. 52. Kolygino, southwestern coast of 

Kamchatka. 

1929. Canis lupus bactrianus Laptev. Opredelitel' mlekopitayushchikh 
Srednei Azii, 1, p. 51. Surkhandar'insk region. 

1930. Canis lupus coreanus. Abe. Joum. Sc. Hiroshima Univers. Zool., 
1, p. 33. Near Seoul, Korea. 

193 1 . Canis lupus hattai. Kishida. Lansania, 3, No. 25, p. 73. Hokkaido, 
Japan. 

1935. Canis lupus rex. Pocock. Proc. Zool. Soc. London, p. 659. 
Hokkaido, Japan. 

1936. Canis lupus dorogostaiskii. Skalon. Izv. Gos. protivochumn. 
inst. Sibirii i Dal'nevo Vostoka, 4, p. 193, Borzya, Eastern 
Trans-Baikaliya. 

1951. Canis lupus hajastanicus. Dahl. Dokl. Akad. nauk Armyansk. 
SSR, 14, No. 3, p. 91. Vicinity of Erevan, Trans-Caucasus. (V.H.). 

Diagnosis 

Body length of adults not less than 90 cm, tail length about half that 
of body. Length of skull more than 200 mm. Arciform line formed by 
anterior border of both nasals, has no process in middle (along line of 
suture between nasal bones). At anterior inner sides of auditory bullae, 

*ln Russian original, misprinted as ciibanenesis — Sci. Ed. 



166 

swellings in form of ridges found on basioccipital. Cingulum on outer 
side of first upper molar narrow and weakly developed, with constric- 
tion or gap in middle. (V.H.). 

Description 

The wolf is a slender, well-proportioned, powerful animal. It has a 
strong trunk with sloping back, with high shoulders and a lower, but 
strong and wide croup. The rib cage is large, deeply descending, the 
abdomen is pulled in and the neck is powerful and muscular. The limbs 
are long and strong, the paws are relatively small and the digits are 
tightly compressed ("in a lump"). 

Head large, heavy, with strong jaws, a long but not pointed, muzzle, 
and broad forehead. Eyes are widely separated and small. Supraorbital 
arches are well developed and therefore the eyes appear to be seated 
125 deeper as if slightly squinting, even somewhat slanting. Ears relatively 
small, triangular in form with pointed tip, directed forward and widely 
separated, and therefore the wolfs head looks especially "broad in the 
forehead". The animal usually carries it [the head] a bit hanging, not 
higher than the level of the back, and appears slightly stooping, with 
its high shoulders. Only the watchful wolf raises its head high."^ 

Tail fairly large, fluffy, hanging down to the tarsal joint. It looks 
as if broken at the base and in the standing and calmly moving wolf, 
it hangs directly downwards. Only during fast galloping does the wolf 
somewhat raise it, and carries it "outstretched" but not higher than the 
back level. In the live animal, the tail moves little and looks very heavy 
(in hunting language it is apdy called a "billet" [chunk of wood]). In 
known psychic states (glad excitement, affection) the wolf wags its tail, 
although not entirely as the dog does. When in fear, it puts its tail 
between its legs as does the dog. Claws black. Eyes yellow. Teats, 5 
pairs. 

The winter fur is very dense and fluffy with short underfur and 
long, quite coarse guard hair. The tail is very densely covered with long 
hairs, which are considerably shorter at the root than along the remain- 
der of its length. In the south, the fur is scarce and coarser, in the 
mid-zone, dense and luxuriant, but fairly coarse, and in the north it is 
longer, denser, more luxuriant and softer. 

'"The wolf is an animal of high psychic [sic] level, and possesses rich mimicry. 
Special study showed that the wolf has ten expressions of the "face" corresponding to 
specific psychic conditions. 



167 




Fig. 44. Wolves, Canis lupus L. Sketch by A.N. Komarov. 



126 



The longest hairs are found on the back, mainly on its anterior part, 
and on the neck. On the shoulder there is usually a region of especially 
long hairs, and on the upper part of the neck the long hairs form a kind 
of crest. The anterior part of the head, including the forehead, is cov- 
ered with short hairs, while on the remaining part, the hairs are longer. 
On the cheeks, the hairs are elongated and form "tufts" — small side- 
whiskers. The limbs, to elbows and a little above the calcaneal joint, 
are covered by short, elastic, closely adjacent hairs. The ears are cov- 
ered with short hairs and strongly project from fur. The summer fiir in 
all latitudes is much more sparse and shorter than that of winter, and 
is coarse and harsh. 

Coloration is monotone, in various parts of the country monotypic, 
and geographically varies comparatively little. Individual variation is 
fairly great, but affects details, while the general color tone and distri- 
bution of colors are constant. Seasonal differences in coloration occur, 
but are not зЬаф, and vary in different parts of the range. In some 
areas, in connection with color fading in winter, the color somewhat 
(sometimes considerably) changes (brightens). Among Soviet wolves, 
melanists, albinos and erythristic are found, but are very rare. Their 
appearance sometimes results from hybridization with domestic dogs. 



168 






''Ш'' 



Fig. 45. Head of adult male wolf. Krasnoyarsk territory. Photograph by 

V.V. Kozlov. 

The wolf of southeastern North America, С /. niger, has diphasic 
coloration — normal and black. 

Sexual differences in color are absent. The coloration of the adult 
Middle Russian wolf in summer fur is composed of a mixture of ocher- 
ous and rusty ocherous tones with light gray. Over the entire skin, 
especially on the surface of the back, there some admixtures of black, 
as a result of the black ends of the guard hairs. The muzzle, approxi- 
mately to the level of the eyes, is pale ocherous-gray, the circumference 
of the lips and the lower parts of the cheeks are white. The space 
between eyes, the forehead, top of head, occiput and the region under 
the eyes and between eyes and ears are gray, only with a light reddish 
film. Around the eyes there are small ocherous rusty fields (rings). The 
backs of the ears are rusty-ocherous with a mixture of blackish-brown, 
and their inner surfaces are covered by dirty white hairs. The chin and 
throat are pure white. The neck is ocherous, somewhat overlaid with 
black (dark ends of hair) on the upper side. 



169 

Long, dark (black) guard hairs along the spine form a well-defined 
black pattern in the form of stripe which is especially bright and broad 
in the anterior part of the back. A quite large amount of hairs with 
black tips is also present on the shoulder blades, and in part on the 
upper part of chest, and the sides on the posterior part of the body. 
However, a dark frosting in the form of obvious "dorsal saddle " is not 
formed. The coloration of the sides of the body and the outer parts of 
limbs is pale dirty ocherous and the inner sides of the limbs are white. 
Belly and inguinal region are white with an ocherous frosting. On the 
anterior side of the fore legs there is usually a shaфly outlined and 
well-defined longitudinal stripe. 

Coloration of the tail corresponds to that of the sides of the body — 
bright dirty ocherous. On the dorsal side, especially on the basal third, 
there is a strong admixture of dark (black, black-brown) hairs; the tip of 
the tail is often black. Hair length in the middle of the back is 60-70 mm. 

The winter fur retains in general the main tones and also distribu- 
tion of the variously colored parts. However, general coloration is 
brighter as a whole in connection with the fact that the underfur is more 
prominent; the tinge of black hairs on this background is more striking, 
appearing as a sort of weak dorsal saddle; ocherous tones appear weaker 
127 and the smoky-gray is more strongly developed. If in some individuals, 
the ocherous tone is clear and quite intensively developed, in some 
others it is weak and the animal looks very light and gray. The length 
of the guard hairs on the shoulders does not usually exceed 90 mm, but 
may reach 1 10—130 mm. 

Age variation in characters of the fur and in coloration is in the 
first year well expressed. Young wolves in their first pelage are cov- 
ered with dense, short (in the middle of the back 20—30 mm), very soft 
"downy" fur of dark-brown or grayish-brown color. This coloration is 
quite evenly distributed on the entire body. The region around the eyes 
and the inner surface of the body are somewhat lighter, while the 
muzzle and the lips, on the contrary, are darker. On the belly the hairs 
are dirty gray with brownish tints and on the chest, between the ante- 
rior limbs, there is a lighter area. The color of the tail, which is covered 
by short hairs, corresponds to that of the trunk. A white tip on the tail 
never occurs. The claws are light. 

This juvenile pelage quickly begins to change and at the end of 
summer the young wolf, still far fi^om attaining the size of the adult (not 
less than two times smaller) has light dirty grayish-ocherous fur, coarse 
and sparse. Reddish tone is not developed and black guard hairs are 



170 






fe - 








Fig. 46. Adult male desert wolf, weight 31 kg. Betpak-Dala desert, 
Kazakhstan. 24 May 1955. Photograph by A. A. Sludskii. 

absent or few. This coloration is quite evenly distributed on the body 
and the light or, on the contrary, dark areas are not differentiated. The 
claws become black. 

From this second pelage, the hairs of which grow greatly by 
autumn, the young ("growing")* wolf passes into the first winter 
pelage. It corresponds to the winter fur of adult animals, but differs in 
the greater monotony of the grayish dirty-ocherous color with less 
development of black and reddish. The color of the wolf in the second 
year ("yearling")**, both in summer and especially in winter, does not 
differ from that of older animals." 

*The Russian word pribylnoi, while literally meaning "growing" or "increasing", 
is best considered the equivalent of "juvenile" — Sci. Ed. 

**Russian, pereyarka — Sci. Ed. 

"Numerous hunters have remarked on age differentiation of wolves on the basis 
of color, as well as have the notes of old authors (Lorents, 1890; Sabaneev, 1877 and 
others) but these are not reliable and invalid (Ognev, 1931; Heptner, 1947; and Heptner 
and Turova, 1951). 



171 

128 The skull of the wolf is characterized by massiveness and generally 
large dimensions. This is the largest form in the family. The facial 
portion, in connection with strongly developed teeth, is relatively long 
and massive; the cranial part of the skull is comparatively small and 
weakly swollen, and is considerably shorter than the facial part, and the 
braincase is relatively small. The nasal bones are long, their posterior 
ends extending to the level of the orbits. Each bone, at the front, is 
notched in an arciform manner in such a way that a common protuber- 
ance is not formed at the line of contact of both bones anteriorly (Fig. 
15). Along the whole length of the line of contact of the nasal bones 
with each other a longitudinal depression (groove) is present. 

The premaxillae gives off a large dorso-posterior process which, 
however, does not reach the frontal bones. The posterior borders of 
nasals and maxillary bones lie approximately on the same level. The 
zygomatic arches are massive- and widely separated, especially poste- 
riorly. The supraorbital processes are large and massive and strongly 
protrude laterally. The facial portion is quite elevated anteriorly and in 
the middle and posterior parts of the nasal bones the profile is some- 
what concave and the forehead abruptly rises upwards, reaching its 
greatest height in the region of the supraorbital processes. 

The frontal region is wide, slightly concave in the middle and 
convex on its margins. Behind the supraorbital processes, the cranium 
is compressed. The sagittal crest is well developed; it bifurcates ante- 
riorly, demarcating the frontal area from the sides and continues to the 
border of the posterior part of supraorbital processes. The occipital 
crest is strongly developed and overhangs the occipital region of the 
skull. The bones of the auditory bulla are of moderate size, thick- 
walled, and their anterior inner parts are not directed to each other, but 
diverge laterally; in this part, at the base of the jugal, are found small 
elongated ridges or swellings. 

The tooth row is very powerful, camassial teeth are massive and 
the canines are strong, relatively short, but with a wide base. 

Age variation of the skull is very great and is reflected mainly in 
the development of the crests, the relative increase in the facial portion, 
increased convexity of the frontal region, and the increased constriction 
of the cranium behind the orbits. 

The wolf cub skull still possessing deciduous teeth is characterized 
by the following characters: facial region of the skull is very short and 
much shorter than the cranial; width of the skull in the region of 
camassial teeth is great; zygomatic arches are situated very close and 



172 

are weak; cranial portion of the skull is relatively large and swollen; 
crests are absent; contour of the braincase region is circular (no pro- 
jections in the upper part of the occiput); no constrictions behind the 
orbits; supraorbital processes not defined; frontal region rises slightly, 
and projections in this part of the skull are absent; auditory bullae are 
relatively large and more rounded; the coronary processes of the lower 
jaw are shaфly curved backwards; and the angular processes are small. 

The skull of the juvenile wolf immediately after tooth replacement 
(in the first autumn of life) has a skull that appears similar to that of 
the adult animal, but differs in the following characters: nasal portion 
is somewhat shorter; skull width in the region of camassial teeth is 
somewhat greater; zygomatic breadth is less; region of the braincase is 
relatively a little longer; crests are absent, and only in the occipital 
region is the posterior part of sagittal crest defined; supraorbital pro- 
cesses are small, short and weakly pointed; constrictions behind the 
supraorbital processes are less, and bones of the auditory bullae are 
relatively larger. 

The skull of the very old wolf, as compared to that of the adult, is 
characterized by a relatively still more elongated facial portion, mas- 
sive and very widely separated zygomatic arches, high and very greatly 
130 developed crests, a wider forehead with greater distance between the 
tips of the supraorbital processes, and shaфer constriction of the brain- 
case behind the supraorbital processes. 

Sexual differences in the skull are manifested only in somewhat 
smaller average dimensions of female skulls. Age changes in their 
skulls are manifested in the same way as in males. 

The length of the intestine of adult Middle Russian wolves (2 
specimens) is 460-575 cm, that of the juveniles 7-8 months old is 390- 
420 cm (3 specimens); the ratio to body length in the former is 4.13 
and 4.62, while in the latter, it is 3.64 and 3.86. The relative weight 
of the heart (Hesse index) varies from 7.32 to 13.07; in young it is 
apparently less than in old (Heptner and Morozova-Turova, 1951). The 
length of the intestine (without the caecum) and the corresponding 
index of two adult males fi-om the extreme north (tundra of Arkhangel'sk 
district — ^Taimyr) are 698 cm and I : 5.3 and 490 cm and I : 4.0. Their 
heart weight is 800 gm and 16.4% (?) and 437 gm and 9.34 % (from 
material of V. Makridin). 

The diploid number of chromosomes is 78, the fundamental num- 
ber, 80 (Matthey, 1949). 



173 





129 



Fig. 47. Skull of wolf, Canis lupus L. Sketch by N.N. Kondakov. 



174 

The dimensions of wolves are subjected to geographic variation. 
Body length of an adult wolf ranges between 105 and 160 cm; tail 
length between 29 and 50 cm (usually from 40 to 50); hind foot length 
is about 220-250 mm; and ear height is about 1 10-190 mm. Height at 
the shoulders is 80-85 cm, and it may reach 100 cm. 

Weight of aduh Middle Russian wolves usually fluctuates between 
32 and 50 kg. Females are not usually so massive as males, of some- 
what smaller dimensions, and their weight is considerably less than 
males. Average weight of females constitutes about 80-85% of average 
male weight. 

Information on the weight of wolves available in the literature, 
especially old hunter's and popular tales, are exaggerated. This is ex- 
plained by the fact that they are mostly based on the determination of 
the weight of particularly large animals "by eye". Recently, in some 
districts, mainly in the middle zone of the European part of the USSR, 
accurate data on the weight of large series of animals appeared (see 
section on "Geographic Variation"). These figures make us deal cau- 
tiously with many of the old data on the average weight of wolves from 
different parts of the range. 

Nevertheless, huge wolves are sometimes found in some places. 
Such animals, however, are very rare. In Middle Russia in general 
form, the maximum weight of wolf is given as 69-79 kg (Ognev, 193 1) 
and 80 kg (Zvorykin, 1939). 

Separate more accurate cases, recently verified, are the following: 
In Saratov district, a wolf weighing 62.4 kg was recorded (Ognev, 
1931); in the forest zone of the European part of the country, 69 kg 
(A.A. Sokolov); in the districts around Moscow, a male weighing 76 
kg is known — this was the largest one among 250 animals killed by the 
famous wolf-hunter V.M. Khartuleri (Heptner ahd Morozova-Turova, 
1951). In the Ukraine, animals of 92 kg (Lugansk district) and 96 kg 
are known (Chernigov district; Komeev, 1950; circumstances under 
which weight of these animals were determined are not known). In the 
Altai, a male weighed 72 kg (Afanas'ev et al., 1953). In the Zoological 
Museum of Moscow University, there is a mounted Middle Russian 
wolf (from F.K. Lorentz), which weighed about 80 kg ("5 poods") 
(V.G. Heptner). 

Information from Vladimir district in recent years indicates that 
here there is a relatively high percentage of large animals. Of 641 
wolves caught during the years 1951-1963 inclusive, 17 animals which 
attracted attention because of their particularly large size were weighed. 



175 

They were caught in Petushinsk, Muromsk, Sobinsk and Suzdal' sk 
regions and had the following weights (kg): Males — 48 (February), 49 
(January), 52 (February), 52 (November), 56 (February), 65 (Decem- 
ber), 68 (January), 70 (March), 76.3 (March), 79 (January); 
Females — 40 (March), 41 (February), 45 (February), 48 (November), 
55 (December), 58 (March), 62 (January). Along with these, three 
aduh wolves (older than two years), but from "the smallest" were 
132 weighed: males 32 and 36 kg (March, April, Gorokhovetsk region) and 
a female 30 kg (March; data of hunting inspection of the district — N.D. 
Sysoev). 

Concerning the Altai animal, it is known that it had "little meat" 
in the stomach, but no other information was given. Sometimes the very 
large weights of individual animals are related to the fact that they 
were supposedly weighed with their stomach quite filled with food; 
recently some have averred that a large wolf can eat 10—15 kg (!) of 
meat. Just now this has been clarified, and information about the vo- 
racity of wolves is greatly exaggerated. Of 1 15 Voronezh wolves, only 
one had 2 kg of meat in the stomach, while all the rest had much lesser 
amounts (Merts, 1953). Of nearly 50 Saratov wolves, none had more 
than 3 kg of food in the stomach (V.G. Heptner). Therefore, the weight 
of some animals under all conditions very greatly exceeds the extremes 
of normal variation (V.H.). 

Systematic Position 

Within the genus Canis, which is considered here in its most narrow 
sense, C. lupus must be considered a more specialized progressive 
form. Its morphological specialization as a predator is, apparently, 
related to its adaptations for hunting large animals. Apparently, with 
this, i.e. with its group way of life in the course of the greater part of 
the year, are also related the progressive, highly developed, psychic 
characters of the wolf (the structure of the pack, the capacity for 
domestication, and so on). Obviously, we must evaluate the systematic 
position of the wolf in the widest inteфretation of the genus (see 
characteristics of the genus and family). (V.H.). 

Geographic Distribution 

Almost all of Eurasia (except the extreme south and southeast) and 
North America (except the extreme south). 



176 




131 Fig. 48. Sole of front foot of a wolf caught in the Sobinsk region of Vladimir 
district. Winter for. Natural size. Live weight of the animal 79 kg. Sketch from 
nature by A. A. Alyapushkin. 



177 

The range in the USSR constitutes a very great part of the species 
range and occupies nearly the entire territory of the country. 

Geographic Range in the Soviet Union 

The wolf is absent only on the Solovetsk islands, Franz- Josef Land, 
Sevemaya Zemlya and the Karagin, Commandor and Shantar islands. 
Sakhalin is also included in the range (Shrenk, 1858; A.M. Nikol'skii, 
1889, Suprunenko, 1890); however, since the beginning of present 
century, it does not live there, and only occurs occasionally as a tran- 
sient, crossing Never sk Strait on ice. Wandering animals do not become 
established (V.G. Voronov and A.G. Voronov). 

Concerning the Kuril islands, information in the literature is in- 
definite and contradictory and, to a considerable degree, erroneous. 
Wolves were recorded on Kunashir and, with doubt, on Iturup 
(Kuznetsov, 1949) and Paramushir (Podkovyrkin, 1950); wolves re- 
corded on Shumishu (Sergeev, 1947) are simply feral dogs (Podkovyrkin, 
1960). According to the most recent data, in the middle of the 60's, no 
wolves were found anywhere on the Kuril islands, and all records 
pertain to feral dogs, which, in particular, are present on Iturup, Urup, 
Shumishu, Kunashir and Paramushir (V.G. Voronov). 

In the north, wolves are met with on Kolguev, Vaigach, and South 
islands of Novaya Zemlya, on White Island, on the coastal islands of 
the western (Nordenskiold island), northern and northeastern Taimyr 
(in particular, Samuil island), on the Lyakhovsk islands and on the New 
Siberian archipelago — on New Siberia and Kotel'n (E.I. Shereshevskii) 
and Wrangel' island. 

At the extreme limits of its range in the north, in particular in the 
northern Taimyr and on the islands to the east of Novaya Zemlya, the 
wolf occurs irregularly and as a transient, not even every year. Thus, 
134 on Wrangel' island only one migration by three animals was known in 
1938 (ManteifeF, 1947). Since migrations here of foxes are known, 
this instance is probable although the width of Delong strait is more 
than 150 km. 

In the Arctic zone, the distribution and intrusions of wolves are 
connected with the distribution and migrations of domestic and wild 
reindeer (see reindeer in vol. I). This applies to individual years as well 
as to considerable periods of time. Thus, wolves which fairly regularly 
inhabited the southern island of Novaya Zemlya have disappeared with 
the disappearance of the wild reindeer from this region and have not 
been met with for a long time there (see "Biology" section). 



179 

To the south of the designated northern boundary, the wolf is 
encountered everywhere, including the extremes of high mountains and 
deserts, though in very different numbers. The wolf is extremely rare, 
and absent in some places over considerable areas only in the taiga, 
which owing to its heavy snows and the relative poverty of food re- 
sources, represent the most unfavorable situation for the animal's life. 
When living conditions change, the wolf also occupies these districts. 
Thus, in the north of Arkhangel'sk district, in connection with clearing 
of forests, the spreading of stock-raising to the north, an increasing 
number of forest wolves are nowadays energetically moving northward 
into the taiga districts which were previously unavailable to them 
(Parovshchikov, 1959). In the last decades, due to felling, road con- 
struction, appearance of villages and similarly due to the disappearance 
in many places of its enemy — the tiger — the wolf occupied a consid- 
erable part of Ussuri taiga where it was previously absent and is 
quickly spreading northward (Abramov, 1954 and others). 

A long time ago wolves were completely destroyed in the Crimea 
and were totally absent there for a long time. They appeared there 
during the Civil War, but they were again eliminated. During the Sec- 
ond World War and the first years thereafter the animals appeared 
again on the peninsula, but quickly disappeared. 

Wolves have migrated across the ice to several islands in the 
northern part of the Caspian Sea, and at the time of seal hunting, 
attracted by the abundance of food (carcasses of killed animals), they 
go far onto the ice of the open sea (from the eastern shore, V.G. 
Heptner). 

Geographic Range outside the Soviet Union 

This comprises Japan (now extinct), the Korean peninsula, China south- 
ward to the Yangtze-kiang, the Mongolian Republic, Tibet, the Hindustan 
[Indian] peninsula to Bengal on the east and at least to Dharwar (about 
latitude 15°30' N. Lat., a little east of Goa) to the south, Baluchistan, 
Afghanistan, Iran, Iraq, the Arabian peninsula, the countries of the 
136 Levant and all of Europe. In Europe the wolf is exterminated in Ireland, 
England, France, Holland, Belgium, Denmark, Switzerland and in all 
of Central Europe. In the German Democratic Republic and even in the 
Federal Republic of Germany during the war and the post-war years, 
the appearance of few vagrant individuals was noted. 



180 




134 Fig. 50. The boundaries of distribution of forest wolf (solid line) and tundra wolf 
[broken line]* in Arkhangel'sk district. Arrows indicate direction of winter 
movements of tundra wolf to the south (after Parovshchikov, 1959). 

In North America, the range of the wolf (reconstructed) occupies 
nearly the entire continent; on the south, to the southern border of the 
Mexican highlands (near 20° N. Lat.), and on the north, the whole 
Arctic archipelago. It also occupies the northern coast of Greenland and 
the western and eastern to 69° N. Lat. (Disko [Island], Godhavn and 
Scoresby Sound). Accordingly, the most northern point of the animal's 
occurrence is located only 700 km from the pole. 

In the south, the range does not include the extreme margins of the 
western coastal parts of the continent, beginning at 41° N. Lat. in the 
south, both in the United States and in Mexico, and the coastal areas 
of eastern Mexico to the Rio Grande. The range also does not include 
the Queen Charlotte islands (it is on Vancouver Island) and the islands 
of the Bering Sea (although they are known to have crossed the ice to 
Saint Lawrence Island). The wolf is exterminated in a considerable part 
of the United States and several races are completely extinct. (V.H.). 



"Not in original Russian — Sci. Ed. 



182 

Geographic Variation 

The huge range and their occurrence in quite different natural condi- 
tions permit us to infer that wolves show considerable geographic 
variation. Authors of the past century have pointed out the formation 
of races in wolves. This is confirmed in our own time by many data. 
However, geographic variation of the wolf is, so far, not satisfactorily 
studied. This applies to the Old World as well as the New where, in 
particular, many improbable subspecies were distinguished. However, 
analyses of several of our populations were made at a good level in 
recent years. These analyses are, however, very few and concerned 
only with craniology. Individual variation in color is quite great, and 
wolf skins are scarce in museums because of their high price. There- 
fore, one may attempt to characterize at the present moment only a few, 
better distinguished races of our fauna, and refer only to what may be 
outlined. This review of the forms and characters given below thus 
bears a preliminary character. 

1. Tundra wolf, C. I. albiis Kerr, 1798 (Syn. turuchanensis, 
kamtschaticus, dybowskii). 

A wolf of large size. Body length of males (23) 118—137 cm, tail 
length 42—52; average weight 40 kg, maximum 49 (1 specimen of 55, 
with the stomach contents). Body length of female wolves (23) is 112— 
136 cm, tail length 41-49 cm, average weight 36.6 kg, maximum 41 kg. 

Greatest length of skull of males (10) 248.5-(M 257.2)-270 mm; 
females 237.5ЧМ 247.9 mm)-256.4 mm. 

Pelage is very long, dense, fluffy and soft. Length of top hairs is 
150—160 mm, guard hairs, 80-150 and underfur, about 70. The usual 
color is very light and gray; underfur has two zones of color: the lower 
is lead-gray, the upper is reddish-gray. 

Concerning the general light color tone, two main types are found, 
connected by intermediate forms: light-gray with reddish shade and 
bluish-gray without this shade. The latter is characteristic of juvenile 
wolves; old animals are lighter in color and have a reddish frosting. 
Moreover, the color of the skin changes noticeably during the course 
of the winter. In November— December, wolves as a group are darker 
than in April— May, when the fur strongly fades under the influence of 
the spring sun. Very light individual wolves are sometimes found in 
May. 

Dimensions and color description apply to wolves of Malozemel'sk 
and Bol'shezemel'sk tundras (Nenets Nats. Okrug, Arkhangel'sk dis- 
trict; Makridin, 1959). 



183 

Dimensions of adult Taimyr wolves (caught in 1961) are as fol- 
lows. Body length of males (1 1), 123-(M 127.7)-H6 cm; of females, 
112-(M121.3bl29 cm; tail length of males, 42-<M 44.4)-48 cm, of 
137 females, 40-(M 42.6)-45 cm; hind foot length of males, 25-(M 26.0)- 
29 cm, of females, 23-(M 25)— 27 cm; ear height of males, 1 1-(M 
.12.7)-13 cm, of females, 1 1-(M 12)-13 cm. 

Greatest length of skull of males (6), 258.0-(M 267.0Ь288.3 mm, 
of females (7), 239.4-(M 251.6)-261.4 mm; condylobasal length of 
skull of males, 238.0-(M 247.6 mm>-*, of females, 221.6-<M 232.3)- 
243.0 mm; zygomatic breadth of males, 134.3-(M 146.9)-160.7 mm, 
of females, 132.5-{M 137.1)-142.2 mm; breadth above canines of 
males, 47.a-(M 51.1)-55.4 mm, of females, 41.4-(M 45.7)-47.3 mm. 

Data on maximum weight of northern wolves were obtained from 
nearly 500 animals caught during 1951-1961, on the Taimyr and west- 
ward to Kanin. The greatest weight was of an old male killed on the 
Taimyr at the mouth of the Dudypta river — ^52 kg. Two animals, killed 
on the tundra east of Kanin and on the Yamal, had weights of 48.8 and 
46.7 kg (without stomach contents) (V. Makridin). 

Found in the tundra zone and the forest tundra of the European and 
Asian parts of the USSR and in Kamchatka. 

Outside the boundaries of the USSR, perhaps in the extreme north 
of the Scandinavian peninsula. 

The belief that the wolves of the extreme north (tundra wolves) are 
the largest race of Old World and are larger than the forest wolves is 
quite widespread. Actually, tundra wolves are, apparently somewhat 
smaller than forest wolves (Makridin, 1959 and others). Statements 
that in tundra wolves "weight reaches 70 kg and sometimes more" 
(Kuznetsov, 1952) as a characteristic feature of the population and the 
subspecies is exaggerated. However, among tundra wolves, specimens 
of more than 50 kg in weight are found. Moreover, it is possible that 
the figures given are somewhat reduced, as they were obtained in a 
locality where intensive hunting of wolves was carried out over the 
course of several years by aircraft (see later). 

As a whole, the tundra wolf is a well-characterized form, and 
noticeably differs from the forms distributed in the forest zone by its 
color and, apparently, its dimensions. At present it is impossible to 
judge whether the wolves in all the area of its distribution are identical 
or not. Concerning color, wolves of the northern Kola Peninsula are on 

*Translator's remark: a figure is missing in original Russian. 



184 

the whole somewhat different, and are more similar to forest wolves 
and wolves of extreme northeastern Siberia. It is not excluded that 
there are also some differences in the average measurements of the 
various tundra populations. They have not been studied, but they do 
not, apparently, exceed the limits of local populations within the bound- 
aries of the subspecies. In the Old World there are no populations of 
almost pure white wolves similar to those living in the extreme north 
of America (C /. bernardi, C. I. manningi and others of the tundrarum 
group). 

This subspecies coincides with the polar assemblage of fiir-bearing 
animals (see beyond). 

2. Middle Russian forest wolf, C. /, lupus Linnaeus, 1758 {syn. 
communis). 

A wolf of very large dimensions, probably maximum within the 
boundaries of our country. 

Color of wolves of this race corresponds to that given above in 
"Description" (p. 166). 

Found in the forest zone of the European part of the country, 
northward to the northern limit of forest, and southward including the 
forest steppe. The exact limits of distribution to the south are not 
known at all; to the west it is to the state boundary and the distribution 
in the east is not established. It probably spreads more or less far into 
Siberia, and possibly, occupies western Siberia. 

Outside the USSR — in Europe except, evidently, the extreme 
south. 

The Middle Russian forest wolf is well differentiated from the 
tundra wolf, although in the north, both forms penetrate into the area 
of occurrence of each other (see "Geographic Distribution"). 

It is usually considered that the weight of adult males is 40-45 kg, 
a yearling, about 35 kg and juveniles about 25 kg. Maximum weight of 
the Middle Russian wolf is considered to be from 69 to 80 kg (see above). 
138 Dimensions of animals from separate regions are as follows: weight 
of adult wolves of northern part of Ryazansk district (south of 
Meshchera — ^a little south of the latitude of the Pra river) varies from 
32 to 55 kg (based on 173 individuals); weight of the largest male was 
56.3 kg (Kozlov, 1955). 

Of 65 wolves killed in the region of Mordovsk preserve (Temnikov), 
only one old male with worn teeth weighed 49 kg. The remaining adult 
males had weights from 40 to 45 kg; adult females, from 30 to 35 and 
only one female weighed 38 kg (Kozhevnikov, 1953). 



185 

The wolves of Voronezh district, i.e. the forest steppe zone, appar- 
ently are somewhat different from Moscow wolves of the true forest 
zone and have the following measurements and weight. Average body 
length, based on 1 54 animals killed in the region of Voronezh preserve, 
was 125 cm in males and 122 cm in females. The majority of animals, 
both old as well as yearlings, had a weight from 30 to 40 kg. Many 
juvenile wolves reached a weight of 23 kg by the end of November. 
Weight of adult (full-grown) males (48) averaged 40.0 kg; weight of 
adult (full-grown) females (31) was 31-42.1 kg, averaging 36.0 kg. 
Weight of yearling male wolves (up to two years old) (8) was 26.6- 
34.0 kg, averaging 32.0; weight of yearling female wolves (24) was 
23.0-32.8 kg, averaging 29.0 kg (weight without stomach contents; 
Merts, 1953). 

Body weight of juvenile and yearling males in Belovezhsk Forest 
varies from 23 to 35 kg; of females — ^from 22 to 29 kg (Table 9*). 

The described race is related to the largest ones and, probably it, 
139 and not the tundra wolf C. I. albus, is the largest form of Old World. 
Beyond that, the figures given show that the average measurements of 
the Middle Russian "forest wolf, usually obtained from especially 
large animals, are often exaggerated. Thus, out of more than 500 ac- 
curately weighed animals, only 3 males had a weight exceeding 50 kg, 
one of these from the Oksk preserve at 56.5 kg (V.V. Kozlov). 

Within the defined region, wolves are not entirely of one type. 
Thus, animals from the extreme west (Belovezhsk Forest) are, as shown, 
somewhat smaller than from Moscow and, in general, those of the 
districts around Moscow, but they do not differ in color. Wolves of 

Table 9. Dimensions and weight (without stomach content) of adult wolves of 
Belovezhsk Forest (Gavrin and Donaurov, 1954)'^ 



Measurements 


Males 


(about 62 


specimens) 


Females (about 58 
min. max. 


specimens) 




min. 


max. 


average 


average 


Body length 


105.0 


140.0 


119.0 


97.0 


124.0 


111.0 


Tail length 


33.0 


50.0 


41.0 


29.0 


45.0 


40.0 


Hind foot length 


20.0 


28.0 


25.0 


21.0 


26.0 


23.0 


Ear height 


11.0 


19.0 


13.0 


11.0 


12.0 


13.0 


Weight 


23.3 


44.8 


34.8 


21.0 


36.0 


29.2 



*ln Russian original, 10, an error — Sci. Ed. 

'^The authors did not give the number of males and females of their series. The 
figures given here are based on the authors' indication of sex ratio in the population they 
studied. 



186 



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187 

Voronezh district are, apparently, not smaller than those of the more 
northern districts, although they are somewhat grayer and lighter in 
color than those of Moscow, the color of which is more reddish. Among 
145 winter skins (mainly from Voronezh preserve) 79% were gray, 
10% light-gray and only about 20% consisted of skins that had "rusty 
and bright-rusty" tone (Merts, 1953). It is probable that the lighter 
color is generally characteristic of wolves of the forest-steppe region. 
This may be the so-called "Central steppe wolf of furriers (see beyond 
p. 195). The Mordovsk wolves correspond to the Moscow ones in color 
and measurements. Difference in measurements in the relatively small 
areas mentioned may not represent geographic variation: in places where 
hunting is very intensive, wolves have little possibility of living to full 
development and attaining maximum size, and the proportion of the 
relatively young animals in the population is large. 

No comparison was made between our wolves and Scandinavian. 
However, judging by the characters of animals from our northwest, 
their assignment to the nominal form is correct. 

3. Siberian forest wolf, С /. altaicus Noak, 1911. 

A large wolf, not less in size than the Middle Russian, C. /. lupus. 

Coloration is light gray, darkened along the middle of the back by 
dark guard hairs. Yellow (ocherous) tones in skin color are absent or 
weakly developed. The fiir is dense, long and soft, but it is slightly 
shorter and not so silky as that of the tundra wolf. Length of guard hairs 
on the withers is 9—1 1 cm. 

Found in Siberia and the Far East except the tundra zone, 
Kamchatka, Trans-Baikaliya and Ussuri territory, and also, probably, 
the western parts of Siberia; northern Kazakhstan. 

Outside the USSR, found in the northern montane taiga of the 
Mongolian People's Republic. 

The systematics of Siberian wolves is not worked out at all, al- 
though wolves in the great territory extending from Urals to the Pacific 
Ocean and from the tundra to the southern mountain ranges are hardly 
identical. According to one view, the area eastward to Lake Baikal is 
inhabited by the nominal form distributed in the European part of the 
Soviet Union (Flerov, 1935), and according to another (Kuznetsov, 
1952), it is the "Siberian forest wolf which is a form different from 
the Middle Russian and occupying all of Siberia except its tundra parts, 
and northern Kazakhstan. 

Until clarification of geographic variation in Siberian wolves, this 
form is conditionally separated. The name given here is also condi- 



188 

tional, it being, apparently, the only one applicable to the Siberian forest 
wolf. Since the western Siberian and northern Kazakhstan wolves are, 
apparently, very close to the Middle Russian ones, then the name C. I. 
altaicus is, above all, applied to the Middle- and East Siberian animals. 

4. Steppe wolf, С /. campestris Dwigubski, 1804 (syn. 
arundinaceus). 

Average dimensions are somewhat less than the Middle Russian 
wolf, C. /. lupus. Pelage is shorter, coarser and sparser. Color of the 
sides is relatively light, gray, on the back rusty-gray or brownish with 
a quite strong admixture of black hairs. 
140 Found in the Pri-Caspian steppes, steppes of the Cis-Caucasus and 
the lower Volga, the steppes of Kazakhstan (except the north?), and the 
steppe region of the European part of the Union. 

Outside the USSR, perhaps in the steppe section of Romania and 
Hungary (in the past). 

Apparently, this is the wolf of the Kazakhstan steppes (except the 
north), intermediate between the northern forest and southern desert 
wolves, which also penetrates into the steppes of the Cis-Caucasus and 
the European part of the Soviet Union. However, it is a form of weakly 
determined characters and distribution. The differences between steppe 
wolves and Middle Russian ones have been mentioned and are accepted 
by all authors, and they obviously exist, although there has been no 
study of this question based upon reliable material. At the present time, 
the wolf is absent or very rare in considerable areas of our European 
steppes. In the northern Ukraine and to the west, also in the steppe 
lives, apparently, not the true, relatively small, steppe wolf, but the 
larger forest or "forest-steppe" form. It is probable that the latter colo- 
nized the westem parts (Odessa, Khersonsk, Nikolaevsk districts), where 
the true steppe wolf was almost completely exterminated during war 
and in the post-war years (Komeev, 1950). 

In the population of steppe wolves, particularly in the Ukraine, few 
very large individuals are encountered (see above). They are mainly 
found in the northern part of the steppe zone. Apparently, these indi- 
viduals intrude from the north (see beyond). 

C. /. campestris, apparently, corresponds to the "Caucasian steppe 
wolf (see later). '^ 



'^In recent years, many references deal with the problem concerning the exist- 
ence in the past in the steppes of Hungary of the very small "reed wolf, 8Ьаф1у 
distinguishable from the large Carpathian forest wolf According to the view of some 



189 

Note: With regard to the type locality selected here (Cis-Caucasian 
steppes) (see synonymy), the following must be taken into consider- 
ation. Dwigubsky states "in desertis ad mare Nigrum et Caspium 
interjacentibus, Kirgisicis, Bucharicis, ad f. Yeniseam, ubi, dantur colore 
sordide flavorubentes" (1804), i.e. "In desert places situated between 
the Black and Caspian seas, in Kirgiz steppes, in Bukhara, and at the 
Yenisei river where dirty-yellowish-reddish forms are encountered", or 
"in the steppes near the Caspian and Black seas; in Kirgiz steppes and 
near the Yenisei river, where red wolves also occur" (1830). 

In all the districts outlined by Dwigubsky, wolves are undoubtedly 
not identical. In accordance with the beginning of the first text given, 
and in connection with the unclear systematic position of wolves in the 
west of our steppes (Ukraine), it is here suggested to restrict the very 
wide definition of Dwigubsky to the mentioned terra typica restricta. 
Dwigubsky [Dvigubskii] moreover stated (1830) that "in my traveling 
in the steppe between the Dnepr and the Don, in 1805 (i.e. after the 
publication of the description. V.H.), I had occasion to see such wolves 
even in the Bakhmutsk district near the Donets." 

The suggestion about the affiliation of the name campestris to the 
jackal (Pocock, 1941) has no scientific basis; in almost all the territory 
mentioned by Dwigubsky, the jackal is absent and was never previ- 
ously found. 

5. Caucasian wolf, C. /. cubanensis Ognev, 1922 (syn. hajastanicus). 

Wolf of medium dimensions (less than Middle Russian). Winter 
coat is poorly furred, coarse, short and quite sparse. Length of guard 
hairs on the withers does not usually exceed 70—75 mm. Coloration is 
dirty, dull gray-ocherous, quite dark due to the more or less evenly 
distributed guard hairs on the skin. Greatest length of skull of males 
240-272 mm, of females 224-251 mm; condylobasal length of skull of 
males is 221—250 mm, of females 209—234 mm; zygomatic breadth of 
skull of males 128—152 mm, of females 11*— 132 mm. 

Found in the Great Caucasus and Trans-Caucasus. 

Outside the USSR it is not known; probably spreads into the con- 
tiguous territories of Iran and Turkey. 



authors, which is, apparently, justified, the "reed wolf is simply the jackal. It is not 
excluded that our steppe wolf described above might have penetrated quite deeply to 
the west. It is not, however, similar to that described by some authors (Nad) under 
the name "reed wolf (Rohrwolf). 

* Error in original Russian text — Sci. Ed. 



190 

141 The distinctiveness of Caucasian wolves is mentioned by a series 

of authors and is, evidently, real. However, the form is very poorly 
studied. Its distribution is also little known. Moreover, in various parts 
of Caucasus, wolves are, apparently, not identical. Thus, on the plains 
of the Trans-Caucasus, the animals are somewhat smaller than those 
inhabiting the northern Caucasus. It is possible that small animals 
belonging to the group of desert wolves {desertorum-^allipes), de- 
scribed from Asia Minor, penetrate here from the south. Wolves of the 
Armenian highlands, described as C. /. hajastanicus, are somewhat 
larger than those of the plains (they correspond, in dimensions, to those 
inhabiting the Kuban Caucasus) and have softer, longer fur (Dal', 
1951; Vereshchagin, 1959). However, their separation as an indepen- 
dent form is insufficiently argued. 

6. Desert wolf, C. /. desertorum Bogdanov, 1882 (syn. 
bactrianus). 

Measurements are considerably less than those of northern forest 
wolves; the smallest form of those inhabiting our country. 

Coloration is light, gray-sandy or yellowish-gray with a few black 
guard hairs on the upper part of the body, mainly along the spine. In 
the area of the occiput and the outer side of the ears, reddish tones are 
developed. Winter fiir is not very luxuriant; quite sparse and coarse. 
The tail is poorly furred. 

Weight reaches 35-40 kg. 

Found in plains areas, including the deserts of Middle Asia and 
southern Kazakhstan north to the middle Emba, and the northern Pri- 
Urals and in the northeast to the region of the Chuya river, Betpak-Dala 
and Pribalkhash'e. 

Outside the USSR, found in Afghanistan and Iran, at least in the 
northern parts of these countries. 

The desert wolf represents a well-differentiated form. Specific, 
precise data on the dimensions of this form have not been published, 
although its small measurements are indicated by all authors. At the 
same time, large individual specimens are encountered in the wolf 
population of Turkmeniya (Heptner, 1956). 

The wolf of the Trans-Caucasian plains and the Mongolian Repub- 
lic has much in common with the desert one. C. I. desertorum is, 
evidently, closely related to the Indian wolf, C. /. pallipes, although this 
form is, apparently, still smaller and also somewhat differentiated by 
coloration and in fur characteristics. The boundary between the geo- 
graphic ranges of the forms desertorum and pallipes is unknown. 



191 

This form, apparently, corresponds to "Middle Asian steppe wolf 
of the Middle Asian bloc of furriers (see beyond). 

7. Tibetan wolf, С /. chanko Gray, 1863 (syn. laniger auct., nee 
laniger Smith). 

Dimensions comparatively small, noticeably smaller than C. /. lupus, 
but slightly larger than C. /. desertorum. 

The coloration is light, whitish-gray with an admixture of darker 
brownish tones on the upper part of the body. Along the spine there is 
a well-defined dark stripe. The winter fiir is soft, long, quite luxuriant, 
with a well-developed undercoat. The tail is densely furred with long 
hairs. 

Weight, apparently, does not exceed 45 kg. 

Found in the Pamirs proper and the Pamir- Alai system and also the 
Tien Shan. 

Outside the USSR, it is found in montane regions of Central Asia, 
Dzhungariya and Tien Shan, across Tibet to Gansu, Sichuan and Yunnan 
in the east and Inner Mongolia in the north. 

This well-defined race corresponds to the "forest (mountain) wolf 
of the Middle Asian fur trade. 

Note: As regards the name and type locality of this form (see 
synonomy) the following must be taken into consideration. "Chinese 
Tatary," from which it was described, is extremely ill-defined. How- 
ever, none of the authors concerned with the nomenclature of wolves 
(Allen, 1938; Pocock, 1941) attempted to specify it. This form has been 
given a very wide distribution in Central Asia "from Chinese Turkestan 
and the Tien Shan through Tibet into Mongolia, northern China, Shanxi, 
142 Sichuan, and apparently, Yunnan; the western Himalayas in Kashmir, 
at least from Chitral to Lahul" (Pocock, 1941). The uniformity of 
wolves in these regions and above all the identity of the animal of the 
low desert of Kashgariya (Chinese Turkestan) and high montane Tibet 
and the Himalayas is very doubtful. Since the long furred wolf is, first 
of all, associated with Tibet, and since the name chanco is employed 
in place of the name laniger to Tibet animals, it is here suggested that 
the terra typica of the form chanco be considered Tibet. 

Within the borders of the Kazakh republic, therefore, the ranges of 
four races of wolf come together. In the north (still 100—130 km south 
of Kustanai, V.G. Heptner) and in the extreme east, the large "Siberian 
[and Middle Russian] forest wolves" (C /. lupus and C. /. altaicus) 
live. In the south is distributed the small, sandy-colored desert wolf, С 
/. desertorum. The entire intervening region is, apparently, occupied by 



192 

a form transitional (intermediate) between them. It evidently follows 
that it should be considered C. /. campestris, the range of which passed 
through the region north of the Caspian Sea westward to the southern 
Russian steppes. In the steppes of Kazakhstan, far beyond the region 
of its usual occurrence, isolated individuals of both the southern and, 
especially the northern forms, wander. Wolves of the northern type 
have been noted at middle Emba (Afanas'ev et al., 1953). In Tien Shan 
the montane form, C. I. chanco lives. There is insufficient basis to 
assume the existence in Kazakhstan of two undescribed subspecies 
(Afanas'ev et al., 1953). 

8. Mongolian wolf, C. /. tschiliensis Matschie, 1967 (syn. 
argunensis, dorogostaiskii). 

Dimensions are not large — ^like С /. desertorum, or somewhat 
larger, but markedly smaller than those of the Siberian forest wolves. 

Coloration is dirty gray, frosted with a weak admixture of ocherous 
color and without bright pale-yellow and chestnut tones. The fur is 
coarse and stiff.''* 

Total body length of males (18), 93-(M 1 15)-158 cm; tail length, 
30-40 cm; hind foot length, 16—24 cm; ear height, 10—14.5 cm; shoul- 
der height, 58-89 cm. Weight, 26-(M 31.3Ь37 kg. Total length of 
females (8), 90-{M 99.6)-109 cm; tail length, 30-40 cm; hind foot 
length, 16-23 cm; ear height 9.5—13 cm; shoulder height, 57—75 cm. 
Weight, 22-(M 27.7)— 30 kg (animals from southern part of Ussuri 
territory; G.F. Bromlei). 

Greatest length of skull of males (5), 226.0-(M 2393.7b253.5; 
condylobasal length of skull (3), 224.0-(M 229.8)-236.5; interorbital 
breadth (5), 38.5-(M 42.0)-45.5; postorbital breadth (5), 38.5-(M 
40.5)-43.5; zygomatic breadth, (5) 114.0-(M 126.7)-139.5 (speci- 
mens from eastern Trans-Baikaliya; Skalon, 1936). 

Found in eastern, southern and southwestern Trans-Baikaliya and 
in the Ussuri territory. 

Outside the USSR, in the Mongolian Republic, except the northern 
montane and taiga parts, in Inner Mongolia and probably, on the Korean 
peninsula. 

The described form is, apparently, quite well characterized — ^it is 
the wolf of the eastern (Central Asian) section of the steppe and desert 



'"Statements that the "eastern (Trans-Baikal) wolves differ by their long tails, in the 
majority of cases curved upwards at the end" (Manteifel' and Larin, 1949) cannot be 
considered seriously. 



193 

zone of Eurasia. Based on its coloration and dimensions, it possesses 
features relating it to C. /. desertorum. Generally, this form is still very 
poorly studied. (Characteristics and distribution based on data of Skalon, 
1936; Loukashkin and Jemakov, 1934; and G.F. Bromlei.) 

Within the USSR, apparently only a narrow strip of the northern 
edge of its range is found; the Siberian forest wolf lives around Lake 
Baikal and, probably, around Chita. 

The presence of this "steppe" form in the forests of Ussuri territory 
is related to wolves from the south having reached there only in the 
present century in connection with the clearing of taiga and the de- 
crease in the number of their antagonist — ^the tiger. Formerly, wolves 
were generally absent. 
143 The nomenclature of this form is unclear. The name employed, 
given all the inadequacies of description is, apparently, concerned with 
it. At present there is no basis for suggesting that the wolves of Trans- 
Baikaliya and northeastern China differ from those from Liaodun*. 
Nevertheless, the name tshiliensis is considered conditional pending a 
solid systematic revision of the wolves of Inner Asia and the Far East. 

9.? Sakhalin wolf, C. /. hattai Kishida, 1931 (syn. rex). 

The characteristics of this proposed form are unclear and its indepen- 
dence cannot be considered established. Its description is unsatisfactory. 
It was described from Hokkaido, where it is now exterminated. It was 
applied to Sakhalin and "perhaps" to the Kuril islands (Нафег, 1945; 
Ellerman and Morrison-Scott, 1951). As shown above (see "Geographic 
Distribution"), the wolf is absent on the Kuril chain, and on Sakhalin, it 
has not been met with since the beginning of the present century. There- 
fore, it is impossible now to decide which wolf lived on Sakhalin. Intrusions 
of the Siberian forest wolf occur from the mainland across Nevel'sk 
Strait on the ice but, apparently, it does not remain. 



Geographic variation in the wolf finds its reflection in the sorting of 
skins into groups, as is usual in the fur trade and this attracted the 
attention of zoologists. The value of these data lies in the fact that they 
are based on the examination of large quantities — ^tens of thousands of 
skins. In some cases, there is a quite exact coincidence of the groups 
with subspecies, while in others, differences in classification are sub- 
stantial. 

*Place in northeastern Xinjiang province — Sci. Ed. 



194 

"Characteristics of winter-killed skins of different groups of wolves 
are as follows: 

Polar group: The entire tundra and forest-tundra belt of the USSR 
and Kamchatka. 

Skin size is very great. In spite of the thin flesh side of the skin, 
large size of skins gives them a considerable weight — one hundred 
skins weigh on average 150 kg. Weight of adult animal reaches 70 kg, 
and sometimes more. Pelage is especially luxuriant, thick, and in par- 
ticular, very soft. Length of guard hairs on the nape of winter skin is 
1 1—13 cm. The abdomen is covered with long dense fur. The flesh side 
of the hide is light and thin. Coloration of the fur is light or dark gray, 
tinged with bluish. Wolves of the Siberian tundra have usually the 
spine and the sides of light, pale-grayish color tinged with blue. The 
dark guard hairs grow mainly on the spine, forming a longitudinal belt; 
on the sides, dark tips of the guard hairs are not well defined. The 
abdomen and lower thorax are whitish. The limbs are almost white. 

In the extreme north of the Far East, wolves have similar color but 
usually mixed with a yellowish tinge. Murmansk wolves have a darker 
color due to the great development and distribution of black tips of the 
guard hairs on the skin. 

Siberian group: All of Siberia and the Far East except the tundra 
regions, and also the northern part of Kazakhstan. 

The group is subdivided into skins of the Siberian forest and steppe 
wolf '5 

Siberian Forest wolf. Skins have the same measurements as those 
of the polar wolf Weight of one hundred hides is 148 kg. Weight of 
144 an adult Siberian wolf reaches 70 kg and even more. Pelage is dense, 
long and soft, but is of lower quality and not so silky as that of the 
polar wolf Length of guard hairs on the nape ranges from 9 to 1 1 cm. 
Flesh side of the skin is compact and quite thick. Coloration of winter 
fiir is usually light gray, darkened along the spine due to dark tips of 
the guard hairs. Yellow tints are either absent or poorly developed. 
Abdomen is whitish. Limbs are gray. 



"Division of the group into subgroups of forests and steppe generally reflects 
the basic types of individual variation in coloration and fur quality. In some cases it 
indicates the migrations (sometimes seasonal) of a certain number of animals of one 
race into a region inhabited by another race. Sometimes, as mentioned above, these 
groups correspond to particular subspecies. 

Data on animal weight, as mentioned above, in the reviewed groups is some- 
times overstated. 



195 

Siberian Steppe wolf. Skin size is as in the forest wolf. Pelage is 
moderately luxuriant, but coarse. Coloration of the spine and the sides 
is gray with clear brownish or reddish tints and with black tips of the 
guard hairs on the spine well developed. Abdomen is reddish or yellow- 
ish. Limbs are brown. 

Kazakhstan group: Kazakhstan, except its northern regions. 

Skins of the Kazakhstan group are also divided into skins of the 
forest and steppe wolf, which are similar to the hides of steppe and 
forest varieties of the Siberian wolf, but differ in their smaller dimen- 
sions and less luxuriant pelage (weight of 100 hides of Kazakhstan 
wolf is 1 12 kg, and length of guard hairs on the withers of forest wolf 
reaches 7 cm). 

Middle Asian group: Middle Asia. 

Skins of this group are divided into skins of forest (more correctly, 
montane) and steppe Middle Asian wolves. 

Forest {montane) Middle Asian wolf. Dimensions are small, and 
weight of 100 hides is not more than 100 kg. Weight of an adult animal 
does not exceed 45 kg. Pelage is quite soft and long, but not particu- 
larly dense. Coloration is light- whitish-gray, slightly darkened on the 
spine by dark guard hairs. Side and abdomen are whitish. 

Steppe Middle Asian wolf. Dimensions are still less than montane 
wolves of Middle Asia. Live weight is usually less than 40 kg. Pelage 
is not very luxuriant, coarse, coloration of spine is light yellowish-gray. 
Abdomen is grayish. 

Central (Russian) group: European part of the USSR, except tun- 
dra regions, Ukraine, Crimea, northern Caucasus, and the lower Volga 
area. They are divided into the forest and steppe central wolves. 

Central forest wolf. Dimensions are large — nearly the same as 
those of the Siberian forest wolf. Weight of 100 hides is 166 kg. 
Weight of live animal is up to 70 kg. Pelage is of average luxuriance 
and softness. Length of guard hairs on the withers is usually not more 
than 9 cm. Coloration of fur in the majority of skins is quite dark, gray, 
usually with brownish tints. The quite dark color of the skin is due to 
there being black guard hairs distributed on the entire middle dorsal 
part of the skin, somewhat darker in the region of the vertebrae. Ab- 
domen is whitish or yellowish. 

Central steppe wolf: Dimensions are slightly less than the forest 
form. Pelage is not very luxuriant, coarse and bristly. Coloration of the 
spine and sides is brownish or reddish-gray. Abdomen is yellowish. 



196 

Caucasian group: Ukraine, Crimea, northern Caucasus, Trans- 
Caucasus, lower Volga area. 

Skins of this group are subdivided into two categories: forest and 
steppe Caucasian wolves. 

Caucasian forest wo//(true montane. V.H.). Dimensions moderate. 
Weight of 100 skins is 146 kg. Weight of live animal reaches 50 kg. 
Pelage is not very luxuriant; sparse, coarse. Length of guard hairs on 
the nape is not more than 7.5 cm. Flesh side of the skin is porous and 
thick. Coloration of the spine and the sides is dirty-gray, usually quite 
dark due to numerous evenly distributed dark guard hairs. Abdomen is 
dirty whitish with yellow tints. 

Caucasian steppe wolf. Dimensions are the same as those of the 

forest wolf. Pelage is short, sparse, and coarse. Fleshy side of the skin 

is thick, porous. Coloration of the spine is usually rusty-brownish or 

145 rusty-gray with dirty tints, and is strongly darkened from dark tips of 

the guard hairs. Sides are lighter in color. Abdomen is dirty-whitish. 

Skins of the polar wolf are considered the most valuable — ^among 
the remaining, the best are skins of the Siberian wolf; skins of the 
Central and Kazakhstan wolves are less valued, and those of Middle 
Asian wolves still less so; the poorest skins are those of Caucasian 
wolves" (Kuznetsov, 1952). 



From parts of the range lying outside the boundaries of our country in 
the Old World, the following forms are recorded. 

1) C. /. signatus Cabrera, 1907 — ^Iberian Peninsula; 2) C. /. deitanus 
Cabrera, 1907 — Spain; 3) С /. italicus Altob., 1921 — ^Italy and Sicily; 
4) C. /. br/flA: Bolkay, 1925 — ^Yugoslavia; 5) C. /. hodophylax Tamm., 
1839 — ^Honshu, Hondo*, exterminated; 6) C. l.pallipes Sykes, 1831 — 
India, Baluchistan, southern Iran, Iraq, ? Asia Minor, northern part of 
the Arabian peninsula; 7) C. /. arabs Pocock, 1934 — southern part of 
the Arabian peninsula. 

The existence of some forms in Europe (two in the Iberian Penin- 
sula) is very doubtful, the remainder are, apparently, real (especially 
С I. pallipes). 



♦Honshu and Hondo are synonyms, the first being the modem, the second the old, 
name of the main Japanese island — Sci. Ed. 



197 

Geographic variation of wolves in the Old World, as regards char- 
acteristics of size and also color is quite regular. The largest wolves are 
northern, in the tundra and forest zones. 

Steppe wolves are somewhat smaller in size, as in wolves of the 
Caucasus, and wolves of the deserts of Middle and Central Asia are 
still smaller. Wolves of this type also inhabit the mountains of Central 
Asia; if some of them are larger than those of desert plains, the differ- 
ence on the average is usually insignificant. The smallest wolves inhabit 
the southern margins of the range — India, Baluchistan, Iraq and the 
Arabian peninsula'*". Wolves of types desertorum, tschiliensis and 
chanco comprise a transitional group from the large northern to 
the small southern ones. Accordingly, Bergman's rule is quite clearly 
demonstrated. 

Wolves with the lightest color are those of the extreme north — ^the 
tundra zone; more southerly ones living in the forest zone are more 
intensely colored, while those of the extreme south — the desert zone, 
have light ocherous "desert" color. Animals of the mountains of Central 
Asia are somewhat more intensely colored than in the desert. Accord- 
ingly, Gloger's law is also demonstrated. 

Geographic variation in New World wolves is well demonstrated. 
Twenty-six well-founded forms are usually accepted (Young and 
Goldman, 1944; Miller and Kellog, 1955; Haltenorth and Trense, 1956; 
Hall and Kelson, 1959)'^. Both a priori and from the published mate- 
rials, it is sufficiently clear that this number is overestimated and the 
number of actually existing and the really differentiated subspecies of 
American wolves is considerably less. Nevertheless, there is no doubt 
that geographic variation of animals on this continent is clearly ex- 
pressed and significant. There is a quite definite geographical trend 
related both to size and color. 

The largest wolves live in the North; towards the south they gradu- 
ally become smaller, having minimal dimensions in the south of the 
range. The largest wolves are in the extreme northwest — ^in Alaska and 
the Mackenize basin {pambasileus, mackenzii, tundrarum). Here ani- 
146 mals attain a weight of almost 80 kg; the known maximum weight is 

'^For the Indian C. /. pallipes, Pocock (1941) recorded a maximum body length of 
100 cm and greatest weight of 24 kg (53 pounds). Weight is, evidently, sometimes more. 

"Since the complete list of these forms is of limited interest to our readers, it is 
not given here. 

The area of all North America is greater than the territory of USSR by only a little 
more than 2 million km^. 



198 

175 pounds — ^79.3 kg; but on average their weight is less, and 45.4 kg 
(100 pounds) is the weight of a large animal. The smallest wolves live 
in the extreme south. Those of Texas and Mexico, C. /. baileyi, and 
still smaller are those inhabiting the extreme southeast, C. /. rufus. 
Weights of the males of this form reach 32-36.5 kg (70-50 pounds), 
but on average the animal weighs 14—16 kg (30-35 pounds). In this 
way, the American wolves follow Bergman's rule more clearly and 
more shaфly than do the wolves of the Old World. 

Coloration of American wolves also changes geographically in a 
regular manner. In the extreme north and northwest live the northern 
tundra wolves. Some races (populations) have a nearly white color (C. 
/. manningi, C. I. bernardi, C. I. tundrarum). Wolves of the forest zone 
have a more saturated coloration, which generally corresponds to the 
coloration of our forest wolves. Steppe, and especially desert forms of 
the south and southwest have light ocherous tones. In the extreme 
southeast of the range, in me sic portions of the country, in the area of 
the lower course of the Mississippi and in Florida, the wolves have a 
very intense color with well-developed ocherous and red tones ("red 
wolf of the Americans). Here, melanists are encountered with greater 
frequency than in any other place, and the coloration of local animals 
is generally dimoфhic'^ In this way, Gloger's rule is sufficiently clearly 
revealed in the described color distribution (V.H.). 

Biology'^ 

Population. Within the boundaries of their range, wolves are very 
unevenly distributed. Their distribution over a territory is conditioned 
by the abundance and the stability of food resources, primarily wild and 
domestic ungulates, and by the possibilities of capturing them, which 
depends in winter on the depth and friability of the snow cover. 

'^American authors usually consider the wolves of the southeast as a separate 
species, С niger Bartrarn, with three subspecies {niger, gregoryi, rufus). This fact is 
hardly well founded. Through the small C. /. lycaon of the southwest, both groups of 
races, "C. lupus'' and "C. niger," merge with each other. Here, the forms of the niger 
group are assigned to C. lupus. 

''Information on the life habits, distribution and abundance of wolves appears in 
a large number of journal notes and articles, and separate books, and in faunistic studies 
and reviews, but only a part of this material represents scientific publications based on 
original and accurately collected data. Therefore, the ecology of wolf cannot be consid- 
ered well studied. Information on the biology of individual populations and subspecies 
is especially scarce. 



199 

On the Kola Peninsula, in the second half of the past century, the 
wolf was very rare, resulting from a decrease in the number of wild 
reindeers and the reduction of domestic reindeer herds of the Saami 
(Laplanders). Wolves at that time were relatively numerous along the 
boundaries with Sweden and Norway, in their eastern regions where 
the wild reindeer was preserved and where domestic reindeer breeding 
was in better conditions (Pleske, 1887). In the Lapland preserve, in 
1929—1938, wolves were totally absent and it was only later that they 
began to bother herds of wild reindeer. In the winter of 1940/41, only 
three wolves lived in the preserve in an area of 1300 kg^ (Semenov- 
Tyan-Shanskii, 1948). 

In the Karelian ASSR, they [wolves] continually inhabited the 
southernmost regions settled by humans. In certain years, wolves were 
numerous in the Pri-Onega and Olonetsk regions, in Trans-Onega and 
in those regions of the republic with little forest, especially along the 
White Sea coast (Stroganov, 1949). The wolf is rare in the Segezhsk 
region, and in the northern regions of the Karelian ASSR with deep 
snow (White Sea, Kemsk and Loukhsk regions), it does not live con- 
tinuously, appearing periodically every 5—10 years. Their appearance is 
connected with the driving of domestic reindeer herds or the migration 
of wild reindeer (Marvin, 1951, 1959). 

In the tundra and forest-tundra of the European part of the USSR, 
wolves are relatively numerous; they accomplish seasonal migrations, 
in summer going onto the tundra and in winter, into the forest-tundra. 
147 In the Mezensk region, wolves are few in summer and numerous in 
winter, since some of the migrants from the Kanin tundra approach 
localities of resident wolves. In summer, only a few pairs remain to 
raise their young on the Koidinsk tundra (Abramovsk coast of Mezensk 
Gulf) and along the northern tributaries of the Nes'. On the Kanin 
peninsula, wolves are encountered the year round. By the beginning of 
winter, the majority of wolves wander into the forest-tundra; some 
migrate to the Mezensk region, but the main mass winters on the 
southern coast of Cheshsk Gulf. To this region also come wolves from 
the Timansk tundra. In summer, not more than 3^ pairs of wolves 
remain in the huge area of the Cheshsk Gulf from Nes' village to the 
lower Pesh (along the Vitasu, Ome, Snope rivers). 

On the Timansk and Malozemel'sk tundras, wolves den along the 
Volong, Travyank, Shchuch'ei, Indige, Belaya, Svetlaya, Kamennaya 
Visk, Vel't', Nerut, and Sul rivers and their tributaries. On the Timansk 
and Malozemel'sk tundras, wolves remain in summer in the Timansk 
range; near the coast, they are exceptionally rare. 



200 

On the Bol'shezemel'sk tundra no wolves are found in the repro- 
ductive period in a coastal strip about 100 km wide, where there are 
few suitable den sites, and in summer, herds of domestic deers are 
pastured along with numerous dogs. Wolves den in the basins of the 
upper courses of the Shapkinaya, Kuk, Lai and Kolva rivers (western 
part of Bol'shezemel'sk tundra), along the upper Adz'va, Bol'shaya 
Rogovaya and Chemaya (central part) rivers, and in the Pai-Khoi range 
and the upper Korotaika, Silovaya and Kara (eastern part) rivers. In 
winter, the majority of wolves migrate from the Bol'shezemel'sk tundra 
to the forest-tundra, from the bend of the Pechora to the upper Usa; 
some move beyond the Urals (Semenov, 1954). 

Wolves are generally fewer on the European tundras than in 
Arkhangel'sk taiga. On the Timansk tundra only 10 wolves were found 
in an area of 10 thousand km^ or 1 wolf per 1000 km^. In autumn, in 
the event of successful reproduction, the number here could be 2.8 
wolves per 1000 km^ (Semenov, 1954). 

In the taiga forests of Arkhangel'sk district, wolves are common in 
the southern and central regions — ^in the basins of the Northern Dvina 
and Onega. Along the valleys of these rivers, and sometimes of the 
Pinega, wolves, singly and in groups, traverse northward. In the 
Priozemii region, from 1947—1952, the number of wolves did not ex- 
ceed 40-45 individuals in autumn (3 wolves per 1000 km^; Semenov, 
1954). In Arkhangel'sk district, the wolf prefers places where the taiga 
is thinned by cutting, the forests are intersected by roads and close to 
villages where there are large areas of fields, floodplains and forest 
cuttings since, in winter, the wolf finds its food only near humans. In 
other northem districts with developing agriculture, the number of wolves 
also increases, and the districts inhabited by them spread northward 
(Semenov, 1954). 

In the Komi ASSR, wolves are rare. In the taiga, they are only 
encountered in the river valleys and lake depressions. They are more 
common in the agricultural landscapes of the southern subzone of taiga, 
and on the tundra. During 12.5 years (1939-1950) in the Komi ASSR 
only 343 wolf skins were tanned. The average number tanned yearly 
from 100 km^ of taiga ranges from to 0.01 ; in regions with agricultural 
and other open lands, from 0.01 to 0.06 skins (Ostroumov, 1953). 

In the Urals, wolves are most numerous in tundra, forest-steppe 
and steppe. Their distribution there is also closely related to human 
settlement. Deep in the taiga, wolves are few or totally absent (Shvarts, 
Pavlinin and Danilov, 1951). 



201 

In Yamalo-Nenetsk Natl. Region, the wolf is met with almost 
everywhere, but dens mainly in the southern tundra beh, in forest- 
tundra, and in the thin forest of the pri-tundra. In winter, it remains in 
the immediate neighborhood of domestic reindeer herds and is, there- 
fore, very rare in the Arctic tundra. In the taiga, it only rarely appears 
in the Krasnosel'kupsk region and in the southern parts of the Purovsk, 
Nadymsk and Shuryshkarsk regions. During a decade (1948-1957), 
1166 wolf skins were tanned from this area (85 to 157 per year) 
(Rakhmanin, 1959). 
148 In westem Siberia, the number of wolves is high in the southern 

tundra, in forest-tundra, southern taiga, birch forest-steppe, and steppe. 
The northern and middle parts of taiga zone are distinguished by low 
numbers of wolves to the west (the regions to the west of the lower and 
middle courses of the Irtysh and the lower Ob'). Wolves only traverse 
the region between the lower Irtysh and middle Ob' in some years. 
Thus, in the Surgutsk region, only 2 wolves in all were caught in 20 
years, and in Vasyugan', only two intrusions were recorded over 12 
years. Breeding by wolves has never been recorded in these regions. 

The somewhat unexpected northeastem maximum in wolf numbers 
in westem Siberia originates in the area with maximal depth of snow 
cover, but this area is notable for its compact snow cover and the 
abundance of wild and domestic reindeer. In the south, regions with 
high wolf density coincide with the places of high number of roe deer, 
regions of developed stock-breeding, and uneven snow cover (Laptev, 
1958). 

Along the Yenisei, in the taiga zone, the wolf is rare everywhere 
until Turukhansk. In the forest-tundra (north of 76° N. Lat.), wolf 
numbers noticeably increase. Places of wolf concentration on the tun- 
dras are restricted to the regions of pastures of domestic reindeer herds 
or stopping places of wild ones (S. Naumov, 1931). Wolves are com- 
mon on the Yenisei delta; periodically they are very abundant along the 
lower Olenek, Yana and Lena [rivers]. In central and southern Yakutiya, 
wolves are few, but in Verkhoyansk territory there are more, and in the 
Kolyma and Chukotsk lands, they are relatively numerous. Along the 
Anadyr, they are common (Ognev, 1926; Sikol'nikov, 1928). In the 
Lyakhovsk islands the wolf, being closely associated with tundra and 
reindeer, is encountered only in summer and in winter migrates away, 
following the deer herds to the mainland (Tugarinov, Smimov and 
Ivanov, 1934). 



202 




Fig. 52. Distribution and relative numbers of wolves in western Siberia (from 

Laptev, 1958): 1 — individual occurrences, 2 — direction of migrations, 3 — area of 

migrations, 4 — ^low population density, 5 — high population density. 

In the taiga zone of middle Siberia, in the Nizhnaya and 
Podkamennaya Tunguska basins, wolves are most numerous in the 
upper Khatanga-Kotui, Vilyuyi, Moiero and the northern tributaries of 
the Nizhnaya Tunguska, where large herds of domestic reindeer graze. 
Along the Podkamennaya Tunguska, to the lower course and southern 
tributaries of the Nizhnaya Tunguska, wolves are scarce because of 
deep, porous snow cover and the small number of wild and domestic 
ungulates. Wolves usually appear here only for a short time (N. Naumov, 
1934). This area of low numbers and nonpermanent habitation of wolves 
149 on the right bank of the Yenisei almost joins with a like minimum in 
wolf numbers in western Siberia (interfluves of the Ob', Taz and 
Yenisei rivers). 

In Lena-Khatanga district, wolves are common along the Olenek 
and along the Muna — a tributary of the Lena. In the Vilyuyi basin the 



203 



too 104 



116 120 




Fig. 53. Distribution and relative numbers of wolves in eastern Siberia (from 

Khudyakov, 1957): 1 — ^wolves numerous, 2 — common, 3 — rare, 4 — transient, 

5 — ^wolves absent. 

wolf is rare, and its numbers increase only above the Arctic Circle 
(along the Olenek). But in the valley of the Vilyuyi itself, they are 
relatively numerous. In central and southern Yakutiya, and also in the 
Kolymo-Indigirsk territory, the wolf is rare even in settled regions, or 
is only found sporadically, but it is common on the tundra and forest- 
tundra. 

In the taiga zone of eastern Siberia, in the Sayans and Altai, the 
wolf is scarce due to deep snow. These predators appear here only in 
those regions where domestic or wild ungulates are common and where 
thirming or reduction* of forests by humans was promoted by reduction 
in porosity and depth of snow cover. In the Atlai and Sayans, wolves 
are few, and in the majority of regions they only appear accidentally. 
In the mountains, in autumn, they follow roe deer into the valleys, 
where the snow is shallower and firmer (Polyakov, 1866; Solov'ev 
1921; Afanas'ev et al., 1953). 

In eastern Siberia, wolves are very abundant along the border with 
the Mongolian Republic; in the southern Pri-Baikal, near Irkutsk, they 



"Misprint in original Russian; svedenie (information) instead of snizhenie — Sci. Ed. 



204 

are common in regions along the railway (Pri-Baikal, Trans-Baikaliya). 
The predators are rare or absent in the northern and a bit of the 
southern taiga in regions of deep snow (Table 11). 

In the taiga of the Far East, the wolf is also rare in all regions of 
deep snow, especially those little opened up by humans. Wolves are 
common in places with abundant wild or domestic ungulates and in 
regions where the taiga has been considerably thinned by logging. In 
Primor'e, the predator is rare — ^along the Iman (Ussuri territory), they 
are encountered principally in its lower course (Zolotarev, 1936). 

In the middle and southern zones of the European part of the 
USSR, including the Pri-Baltic, wolves are rare. Their number grew 
150 during war time, 1941—1945, but as a result of intensive control, at the 
present time their numbers are again reduced. In Belorussia, the wolf 
is met with everywhere. In 1932—1939 it was most numerous in Minsk 
district, and later in Gomel' sk, Mogilevsk and Vitebsk districts (Table 
12). 

In the central chernozem districts, not long ago wolves were quite 
common. Relatively large numbers were noted in 1951—1953 in 
Voronezh and Belgorodsk districts (Barabash-Nikiforov, 195.7). 

In southern Bashkiriya at the edge of the upland-forest landscape 
in the forest-steppe, wolves were few in the 30's. About 1950, they 
were quite often encountered, but in the upland broad-leaved forests, 
they became scarce in winter (Kirikov, 1952). Wolf numbers here are 
sharply reduced. In Volzhsk-Kamsk territory, from 1950 to 1965 wolf 
numbers sharply decreased. In Tatariya during the five years from 
1960-1964 the average annual production of skins was 170 (140- 
186, V.A. Popov). 

In the Ukraine, the wolves multiplied during the war years, 1941— 
1945. In 1947-1949 their numbers in the entire Ukraine reached 
about 7000. Population density was low in those places where vil- 
lages were closely situated (Komeev, 1950). After 1945, wolves were 
fi-equent in the southern part of Sumsk district (especially in more 
wooded regions). In Chemigovsk district, which is very suitable for 
wolf habitation, their high numbers were retained the entire time and 
in 1938, 110 animals were caught here. After the war, especially 
many appeared in Chernigov Poles'e; fewer in the forest-steppe and 
southern regions. In Kievsk district, the predators became numerous 
in its northern, forested sections. They were occasionally captured in 
the regions adjoining the Poltavsk and Chemigovsk districts; in Kievsk 
district, wolves were noted in a series of southern regions. In 



205 



Table П. Manufacture of wolf skins from eastern Siberia (Khudyakov, 1937) 



~ ....„^^^^Years 


1932 


1933 


1934 


1935 


Average, 


Regions ^~"~~~~~~...,>.^ 










% 


Main Western 


398 


315 


309 


292 


11.0 


Eastern 


1478 


1175 


905 


1369 


42.4 


Remote taigas 


16 


7 


20 


17 


0.6 


Northern 


38 


33 


46 


89 


1.7 


Buryat ASSR 


1636 


1186 


1266 


1239 


44.3 



Table 12. Average annual output of wolf skins per 1000 hectares in 
Belorussia (Serzhanin, 1955, with modifications) 



151 



^*\^ Years, data 


1932- 


-1939 


1948-1952 


^^^^ 


Average 


Fluctuation 


Average 


Districts ^\^^ 


by di.strict 


m various 
regions 




Brestsk 


— 


— 


0.5 


Vitebsk 


0.1 


0.8-0.02 


0.3 


Grodnensk 


— 


— 


0.5 


Gomel 'sk 


0.2 


0.8-0.03 


0.3 


Minsk 


0.3 


0.8-0.03 


0.4 


Mogilevsk 


0.1 


0.3-0.03 


0.4 


Molodechnensk 


— 


— 


0.4 



Zhitomirsk district, wolves were restricted to the northern regions, 
along the border with Belorussia (Komeev, 1950). 

In the western districts of the Ukraine, wolf numbers are generally 
not great. They are higher in western Poles 'e, in Rovensk and Volynsk 
districts. There are many wolves in the montane regions of former 
Stanislavsk district. In the Trans-Carpathians and in L'vovsk district, 
the wolf is rare; in Ternopol' sk district, it does not live permanently, 
appearing periodically from the southern regions of Rovensk and the 
northern regions of Stanislavsk districts (Tatarinov, 1956). In L'vovsk 
district, wolf litters were noted only in the Kamensko-Bugsk, Sokal'sk 
and Peremyshlyansk regions. In the Trans-Caфathian area, they are 
relatively common in all montane and low-lying regions (except rocky 
places in the high mountains). Stray wolves were met with in the 
Vinogradovsk, Uzhgorodsk, Perechinsk and Mukachevsk regions. In 
Drogobychsk district, wolves are common in montane regions; in 1949 
litters were encountered in the Turkovsk and Podbuzsk regions, and 
strays in the Strelkovsk and Dorogobyshsk regions. 



206 

In Stanislavsk district, wolves occupy permanently the southern 
and southwestern montane region, and intrude on the plains. In 
Chemovitsk district, wolves occur permanently in five southwestern 
regions bordering Romania, and also in Sakyrsk and Kal'menetsk re- 
gions, where they often come from Moldavia. 

In Kamenets-Podol'sk district, wolves are established in the north- 
em regions, but animals are scarce here. In Vinnitsk district, wolves 
are regularly noted only in two southern regions; they penetrate hither 
from Moldavia, where there are many wolves. In Kirovogradsk district 
wolves are more or less constantly present in the Pri-Dneprovsk re- 
gions adjacent to Kiev district. In Poltavsk district, wolves live in the 
eastern regions (near Sumsk district), and penetrate along the forested 
massif into Mirgorodsk region; they were also found nearer to Kiev 
district. 

In Khar'kov district after the war, wolves propagated more in the 
central regions. Very few litters were found in the Volchansk region, 
and in the Kupyansk region vagrant wolves were recorded. In the 
Lugansk district, wolves were present in 3 1 out of 34 regions; they 
were especially numerous in the northeastern regions. In six of these, 
in 1949, 150 wolves were captured. In Donetsk district, wolves are rare 
and noted only in the south. In Dnepropetrovsk district after war, wolves 
lived permanently only in the extreme eastern regions. In Zaporozhsk 
district, wolves live mainly in places near the Dnepr floodlands, and 
appear occasionally in the Berdyansk and Pri-Azovsk regions. 

In IChersonsk district until 1948—1949, the Tsyurupinsk and 
Skadovsk regions were densely inhabited by wolves; on the right bank 
they were totally absent; after 1948, they appeared in the Khersonsk, 
Berislavsk and Novovorontsovsk regions. In Nikolaevsk district, wolves 
live permanently in the Shirokolanovsk region; strays appear in the 
western regions. In Odessa district, wolves are common in the western 
regions. In 1949—1950, strays were also observed in the central re- 
gions. In the Izmail'sk region, wolves live permanently in the regions 
adjacent to the Moldavian SSR. In the floodlands of the Dnestr, wolves 
are few (Komeev, 1950). In the Crimea, wolves were exterminated, but 
during the years of the Patriotic [2nd World] War, they appeared again 
on the peninsula; they were soon destroyed. 

In the Caucasus, the wolf is distributed everywhere, though not 
evenly. According to the data on tanned skins, the greatest number of 
individuals per unit area was captured in the western Cis-Caucasus and 
eastern Trans-Caucasus. In the 1930's of this century in Azerbaidzhan, 



207 

were caught per 1000 km^, 1 1.6 wolves; in Armenia — ^9.9, in the north- 
em Caucasus, 6.7; in Dagestan, 6.7; and in Georgia, 0.8 wolves. The 
lower Terek, Sulak, Kuma and the coast of Kyzylagachsk Gulf were 
the least densely populated by the wolf. In alder bogs of Kokhida, wolf 
is common in the lower Gagid, Okum and Picher rivers. In montane 
regions of the Caucasus, wolf numbers are high in the Caucasian, 
152 Borzhomsk, Zakatal'sk and Lagodekhsk preserves (Vereshchagin, 1959). 
In the Trans-Caucasus, wolves are everywhere, but along the wooded 
Black Sea coast they are so rare that they were never seen by the 
majority of inhabitants (Satunin, 1915). 

In Kazakhstan, the wolf is encountered nearly everywhere. Their 
numbers are highest in the west, in several regions of Aktyubinsk and 
Kustanaisk districts, in the valleys of the Syr-Dar'ya, Chu, and Hi 
rivers, and in the mountains and foothills of the Zailiisk and Dzhungarsk 
Alatau, near Lake Kurgal'dzhin, in the Alakul'sk depression, around 
Zaisan Lake and in several other places. In the northern forest-steppe 
regions, wolves are scarcer. However, in the former Kustanaisk district 
of the former Turgaisk governance, their number in beginning of the 
twenties was extremely high (see beyond, V.G. Heptner). In clay deserts 
and semi-deserts, where there is neither water, wild ungulates or live- 
stock, these predators are absent or appear there only from time to time. 
There are also no wolves in some of the montane taiga regions of the 
southern Altai, for example, in the Zyryanovsk region, where snow 
cover is deep and porous (Afanas'ev et al., 1953). The general number 
of wolves in Kazakhstan is estimated at approximately 25—30 thou- 
sands. About 10 thousand wolves are exterminated annually in 
Kazakhstan; i.e. a little more than one third of the population (Afanas'ev, 
1955). 

In Kirgiziya, wolves are encountered everywhere. They are most 
numerous in the regions of large high montane pastures-elevated 
watersheds, since in these places not only many livestock are found but 
also great marmot colonies were preserved, and in the neighboring 
ranges, large numbers of wild ungulates (arkhar sheep and wild goats). 
Each year in Kirgiziya, prior to the Great Patriotic War, up to 400— 
600 wolf skins were tanned (Kuznetsov, 1948). 

In Uzbekistan, there are few wolves. In Kashkadar'insk district 
they are rare; they are rarely met with in desert regions, even in winter; 
in montane regions, they are numerous (Meklenburtsev, 1958). 

In the Turkmen SSR, the distribution of wolves is restricted by the 
presence of water sources and the abundance and also availability of 



208 



60 65 70 76 80 85 



125 о 125250376km 




Fig. 54. Relative number of wolves in various parts of Kazakhstan (from 

Afanas'ev et al., 1953): 1 — general distribution, 2 — districts where wolves are 

especially numerous. 

food. In Badkhyz (southern Turkmeniya), there are more wolves, ap- 
parently, than in other regions of Turkmeniya, which is explained by 
the abundance of wild and domestic ungulates. The especially large 
number of wolves between the Tedzhen and Murgab has already been 
shown by Radde and Walter (1889). Along the Kushka river wolves 
153 frequently hunt near the most remote villages. Along the Egrigek river, 
there are more wolves than along the Kushka; although here there are 
no human settlements and livestock, dzheiran gazelles [Gazella 
subgutterosd] are numerous, which come to the river for drinking and 
there are also abundant watering sites and convenient places for dens 
and diurnal lairs. In Gyaz'-Gyadyk, wolves are common in the whole 
region. There are many wolves along the Tedzhen river. In the desert 
part of Badkhyz, the wolf is everywhere, but is unevenly distributed, 
being concentrated by wells, in grazing sites of sheep flocks and near 
waterholes, especially those situated not closer than 100-200 m from 
domiciles (earthen huts). Six to seven wolves on average are associated 
with one waterhole. In winter, wolves are evenly distributed in Badkhyz, 
because of the reduced need to drink and the wider pasturing of the 
sheep flocks (Heptner, 1956). The number of wolves in western 
Turkmeniya is high, but there distribution is uneven. 

It is difficult to estimate, even approximately, the general number 
of wolves in USSR, not only because of the absence of data, but also 



209 

because of uninterrupted changes in their numbers. In the 1930's, their 
number in our country was estimated by a series of authors to be 60- 
80 thousands, a figure which was, apparently, an underestimate. At 
nearly the same time, S.A. Buturlin evaluated the wolf population as 
being not less than 100,000 individuals, and the yearly production as 
47,000. After the Patriotic War of 1941-1945, the number of wolves 
markedly increased, particularly in the regions which suffered from the 
German invasion. After restoration of the destruction caused thereby, 
as a result of augmented wolf control and the application of a series of 
very effective means for their extermination (shooting from aeroplanes 
and airosled*, etc.), the number of wolves was reduced in the 1960's 
by not less than 3—5 times and in places, more. In a series of districts 
they disappeared entirely. 

One may establish several general laws: 

1) The highest numbers of wolves are in so-called "open" land- 
scapes — southern tundra, forest-tundra, forest-steppe, steppe and several 
desert regions. 

2) Within these zones, the distribution of the predators and their 
mode of life, in particular nomadism, depends upon the allocation of the 
chief foods — ^wild and domestic ungulates. 

3) In the forest zone, especially in the taiga, the most important 
obstacle for wolves reveals itself to be deep, porous and prolonged 
snow cover, which shaфly limits the possibility of movement and cap- 
turing food. The rarity or absence of wolves in the deep taiga regions, 
which is a consequence of this situation, was already established by 
A.F. Middendorf (1869)** and later confirmed by all investigators. 
However, a clear coincidence between the snow cover depth and the 
abundance of wolves cannot be established, since the relation here is 
more complicated; in actuality, the influence of snow on the number of 
wolves acts through a complicated system of biological relationships 
and essentially changes through the direct and indirect effects of human 
activities. 

The construction of roads and paths increases the possibility of wolf 
movements, while the added complexity of the landscape and the appear- 
ance of farming and animal husbandry in these regions enriches the potential 
food reserves, especially in winter time. This explains the direct relation 
that is manifested between the number of wolves and the agricultural 
development of the territory in little-developed taiga regions. 

*Foreranner of snowmobile — Sci. Ed. 
**Not in Lit. Cit. — Sci. Ed. 



210 

In many poorly inhabited regions of taiga, there are no permanent 
wolf populations and the animals only rarely intrude into these places 
by chance. In such unfavorable and almost uninhabitable territories, 
wolves penetrate mainly along the river valleys with their rich and 
complex biocenoses and their greater degree of development of places 
suitable for humans. 

4) In the desert-steppe regions, the presence of watering places 
serves as the important factor limiting the number and distribution of 
wolves, being quite necessary for the predators, especially in the hot 
154 time of the year. There, concentration of the animals is noticeable near 
wells and in river valleys, where they find, not only the necessary 
water, but also abundant food smce in such places both wild and 
domestic hoofed animals are concentrated. 

Habitat. Places of diurnal rest, where dens are also located during 
the reproductive season, are usually especially well protected and are 
characteristically near watering places. The hunting territories of wolves 
are various, and are restricted only to the possibilities of capturing food. 

On the tundra, places of diurnal rest of wolves are restricted mainly 
to river valleys and thickets in dry plateau sections; in the forest-tundra, 
forest clearings and osier-beds and thickets of dwarf arctic birch serve 
this puфOse. In the Yamal and Bol'shezemel'sk tundras, wolves con- 
struct dens, mainly among bushes in valleys and more rarely in the dry 
meadows of slopes; they use flood terraces and dry osier-bed thickets 
along drainage divides. Of 1 1 biotopes in river valleys, wolves use 
three, and on interfluves of dry plateaus, of 17 biotopes, they are 
distributed in only one (dry shrubs along the divides; Sdobnikov, 1937). 
Wolves colonize sea coasts only residually*, where they often feed on 
marine flotsam (Sokolov, 1951). 

In the forest zone, wolves avoid monotonous dense forests. In the 
taiga of West Siberia, they live chiefly in the river valleys (floodplains) 
(Laptev, 1958). In the Irkutsk district, they prefer places where small 
sections of forest alternate with fields, and in winter they use roads and 
paths laid out by humans widely (Timofeev, 1941). In the Karelian 
ASSR, wolves live mainly near populated places, in cut-over forests 
and shrubs adjoining fields. In winter time, they not only approach 
populated places, but often enter them (Marvin, 1959). In Byelorussia, 
avoiding the large forest massifs, they prefer small dense plantations 
with thickets (Serzhanin, 1955). 

*The Russian word gushcha (sediment, dregs) is used here — Sci. Ed. 



211 

In heavily settled regions, wolves, though living in the immediate 
neighborhood of villages, stay in especially secluded places: forest 
islands, bogs, remote ravines and gorges, and dense growths of shrubs; 
at the same time, they often live in vacant weedy field, and sometimes 
in crop fields (Mertts, 1953). 

In the steppes, wolves spend the day, and in the reproduction 
season construct dens, in growths of shrubs and the remote ravines and 
gorges overgrown by weeds, among tugais [riparian thickets] in the 
valleys of rivers and streams, in the old weedy fallow fields and in crop 
fields. In western Kazakhstan, wolves are common in summer in reeds 
by water bodies, in ravines and sand hills, if there are waterholes there. 
In winter, they concentrate near livestock pastures, in reeds along the 
banks and by large lakes, and also near villages, where they feed on 
carrion and hunt dogs. In northern Kazakhstan, wolves live in narrow 
gorges, in vacant lands among weeds, and in forest clearings choosing 
elevated, drier places. In winter they spend the night there and gener- 
ally remain close to human settlements which they approach by night, 
while during the day they hide in weeds or reeds by lakes (Afanas'ev 
et al., 1953). In the Kustanaisk intact steppes, they find shelter in 
growths of dwarf almond {Amygdalus nana) and steppe cherry (V.G. 
Heptner). 

In the deserts and. semi-deserts of central Kazakhstan, wolves are 
restricted in the warm time of year to melkosopochniks near springs 
and streams, in deep sayakhs* (ravines and valleys) if water sources 
are found in their bottoms, in weeds by water bodies and in forest 
islands. In winter, they concentrate in regions of livestock husbandry, 
near rivers and lakes, or they follow the herds of saigas and dzheiran 
[Gazella subgutterosa] (in Betpak-Dala). 

In southern Kazakhstan, especially numerous are wolves living in 
the river valleys (Aksu, Karatal, Hi, Chu, Talas, Syr-Dar'ya). In sum- 
mer, they are found in the tugais, reeds and rose willows at the lake 
channels, and rarely in the stabilized sand hills near water. Here, many 
wild pigs, roe deer, hares, pheasants and waterfowl are found year- 
155 round; here also domestic cattle graze. In winter, the wolves move, by 
night, from the river valleys to adjacent regions where domestic cattle 
and dzheiran winter, but at daybreak they again return to the valleys. 
In the sands, wolves only occur near waterholes, constructing dens in 
saxaul forests or in dense growths of shrubs (Afanas'ev et al., 1953). 

*Foreign (Turkic?) word — Sci. Ed. 



212 

The distribution of wolves among the biotopes in other desert re- 
gions of Middle Asia is the same. In Turkmeniya, the wolf is widely 
distributed, but is obviously attracted to human settlements and places 
where domestic livestock graze. It is encountered high in the mountains 
(2000 m above sea level and higher), in sandy and clayey deserts and 
cultivated landscapes. In intense heat, wolves are almost absent deep 
in the desert, or they are rare and are restricted to only the few wells 
which are used by people where there are sheep flocks and also water 
remaining from the drinking places of the herds. The animals occur 
mainly along the river valleys near the watering places of wild ungu- 
lates (mainly dzheiran) and domestic cattle. In winter, when the wild 
ungulates and herds are distributed throughout the whole desert, wolf 
distribution changes accordingly (V.G. Heptner). 

In the mountains, the wolf ascends to a height of 3 and 4 thousand 
m above sea level (Pamir). In the mountains, the thick and very snowy 
cover favors wolves because they, in following ungulates, stick to less 
snowy slopes and those blown free. 

In the northern Caucasus, wolves prefer inhabited places, and in 
remote places, are met with only where there are many wild ungulates. 
The life of the wolf here is closely connected to the domestic animal 
herds, especially sheep flocks, in the following of which the predators 
ascend the mountains in summer and in winter, descend (Satunin, 1915). 
In the Caucasian preserve, 30% of the incidence of wolf encounters 
were in the alpine zone, in dark coniferous forests, 32%, and in the 
broad-leaved forests, 38% (Teplov, 1938). 

In Armenia, the wolf inhabits the dry subtropics, semidesert, mon- 
tane and meadow steppes, sub-alpine and alpine meadows, and also 
encountered in barren mountains, cliffs and talus, and do not avoid 
cultivated lands. Within the boundaries of Armenia, the wolf is met 
with at heights of 560-3800 m above sea level. In summer, a greater 
portion of the wolves hold to high mountains, in places where domestic 
cattle graze, and are in small number in the desert and barren mountain 
zones. By winter, wolves descend to the valleys and only a few of them 
remain in the mountains near herds of wild ungulates (Dal', 1954). 

In the mountains of Kazakhstan and Kirgiziya, wolves remain in 
summer in alpine meadows, abounding in marmots and wild ungulates, 
and where the domestic animals pasture at this time. In the upper 
reaches of montane rivers (at heights of 3000-3500 m above sea level) 
wolves also raise their cubs at this time. After the development of snow 
cover in the mountains, following the wild and domestic ungulates, they 



213 

descend to the foothills and valleys; only a small number of them 
remain near the herds of wild, and sometimes domestic, ungulates, 
remaining in the mountains in winter in the heat of the sun in sunny and 
wind-blown areas (Afanas'ev et al., 1953). 

In Semirech'e there are, apparently, two populations of wolves; 
one lives in the mountains and descends in winter together with the 
ungulates, wintering in the foothills, while the other — the plains popu- 
lation — occupies in summer impassable thickets of sea buckthorn, 
tamarisk, and other shrubs in the tugais. Both populations are associ- 
ated with flocks of sheep and wild ungulates (Shnitnikov, 1936). 

Food. The wolf is a predator, the main food of which consists of 
mammals of moderate and large size. Almost everywhere it is associ- 
ated with the ungulates comprising its main food. The quantity and 
accessibility of it determines the predators' numbers. Upon the life 
characteristics of the ungulates depends also the way of life of wolves, 
differing in different regions because, first of all, the species composi- 
tion and biology of the wild ungulates or the methods of keeping domestic 
animals in these regions are not the same. If not fundamental, then of 
significant importance in the nutrition of wolves are animals of medium 
156 and small size — marmots, hares, badgers, foxes, polecats and others. In 
many places, wolves successfully hunt dogs, and, in the regions where 
raccoon dogs are acclimatized, they are destroyed in great numbers by 
them. Among small mammals, wolves catch ground squirrels, mice, 
hamsters, voles and other rodents, and also insectivores. In the agricul- 
tural forest-steppe and steppe regions, wolves eat small rodents in the 
fields. They successfully hunt waterfowl, especially during the time of 
their molt. Large birds, mainly egg clutches and young, suffer from 
wolves. Wolves destroy many domestic and wild geese. Reptiles (liz- 
ards and snakes), frogs, rarely toads, and also large insects are eaten 
by wolves when other foods are insufficient. In times of hunger the 
predators readily eat carrion, visiting cattle burial grounds, slaughter- 
houses, tallow-presses or especially situated lures. If such places of 
carrion deposition are permanent, they can determine the winter route 
of wolf packs. 

In many, if not all, regions, wolves are partial to plant as well as 
animal food. They willingly eat berries of mountain ash, lily of the 
valley {Convallaria majalis), bilberries, blueberries and cowberry (in 
the forest zone), nightshade (Solanum nigrum), fruits of apple, pear 
and others (in the south). In summer, they readily visit melon fields, eat 
watermelon, muskmelons and frequently cause great damage, not only 



214 

by eating the fruits, but many others are spoiled. They frequently eat 
various grain crops, and in the Pri-Ural steppes, the delicate and sweet 
shoots of reeds. 

Cannibalism is not rare among wolves. In times of hunger in win- 
ter, the pack often attacks weak or injured animals. They may attack 
a male, which in its struggle for a female, becomes severely wounded. 
In captivity, cannibalism is noted during the transition of young wolves 
from meat to milk or vegetable foods. The stronger cubs attacked and 
ate the weak (Barabash-Nikiforov, 1957). Hungry wolves fight fiercely 
for food, and frequently kill the weaker ones, which are afterward 
almost always consumed. Cases are described when wolves killed and 
ate the wounded or the dead bodies of their kin (Makridin, 1959). 
Therefore, these animals are generally not fastidious about food, but 
when possible, they eat only the best food; this is particularly related 
to the summer time and concerns plant foods, the need for which is not 
so large. 

The wolf is an animal of great endurance. It is able, without losing 
strength and running speed, to survive prolonged hunger, for a week or 
more. But upon succeeding in a hunt, it is very voracious and, accord- 
ing to some authors, can eat immediately a large quantity of food — ^up 
to 25 kg, as if laying in a store (ManteifeP and Larin, 1949; Timofeev, 
1949; Semenov, 1954). It was noticed that a litter of 7-10 wolves 
completely consumed an entire horse carcass during one night (Sokolov, 
1951). In the lower Hi river (Kazakhstan) a pair of wolves ate a roe 
deer weighing 25—30 kg or a young wild swine of 30-40 kg at one time 
(Afanas'ev et al., 1953). In Badkhyz (Turkmeniya) a wolf was ob- 
served to devour a young arkhar sheep of about 10 kg in weight 
(Heptner, 1956). However, these amounts do not characterize the quan- 
tity of food actually eaten at one time. A part of this food is usually 
pulled apart and hidden away, especially when there is plenty of food. 
At the same time, animals torn apart by wolves are often further eaten 
by hyaenas, jackals and especially vultures. In Badkhyz, a skeleton of 
a donkey which was torn apart by a pair of wolves at the start of night, 
was found in the morning completely cleaned of meat (Heptner, 1956). 
More than 1.5—2 kg of food can be rarely found at one time in the 
stomachs of wolves (Novikov, 1956). According to the accurate data 
of P.A. Martts (1953), a wolf eats no more than 3 kg of food at one 
sitting and any excess amounts are quickly regurgitated. The relatively 
small amounts of meat eaten by a wolf at one time is also to be 
understood by the figures given above on the weight of wolves (p. 174). 



215 

The foods of wolves vary substantively in the various seasons, and 
these changes are associated with changes in the life habits of the 
157 predators — their transition from a settled life in the warm time of year 
to a nomadic way of life in winter. In summer, various foods are 
available for wolves, and its quantity is maximal in this period. There- 
fore, summer feeding of wolves is diverse. The principal share in various 
regions is occupied by different foods, but mainly by animals of me- 
dium and small size. The significance of ungulates is reduced in this 
period, although wolves still hunt them. Thus, in the stomach of a 
young wolf killed on 3 July 1944, in the Bashkir preserve, remains of 
a young maral [Cervus elaphus], a mole's foot and 2 nestling sparrows 
were found; and in the stomach of a juvenile wolf caught on 17 August 
1941, 16 lizards were found. In the valley of the Ural river, wolves in 
summer frequently visit melon fields and eat watermelons, and in the 
Shaitantau [mountains] during a year of steppe-cherry harvest, its stones 
were found in the majority of the wolves' feces (Kirikov, 1952). At the 
end of summer and the beginning of autumn wolves hunt pastured 
livestock and wild ungulates, catch hares, dig up muskrat burrows and 
houses, catch the small animals and birds, and around water bodies, 
successfully obtain molting waterfowl. 

After snow accumulation, feeding worsens. At this time, the prin- 
cipal attention of wolves is directed to ungulates. In the time of greatest 
hunger, wolves approach populated areas, hunt dogs, attack livestock in 
their sheds even by day, and willingly make use of cattle burial grounds 
and carrion. 

In winter, wolves travel along roads and unwillingly turn into the 
snow upon the appearance of not only a single sledge but even a series. 
At this time, wolves also attack elk*. However, the attack by an indi- 
vidual wolf of an adult elk, especially in more or less deep snow, often 
ends in the death of the wolf. For instance, in the winter of 
1952/1953 in Verkhne-Toemsk region of Arkhangel'sk district, two 
wolves were found killed by elks (Semenov, 1954). An attack by a 
pack usually is successful. In March— April, in the forest zone on crusted 
snow, wolves rarely appear near the villages and mainly travel through 
the forests, hunting roe deer, elk and [reinjdeer on the snow crust. 

Early spring (after the crusted snow) is the time of greatest hunger, 
when wolves are very harmful to animal husbandry (especially in the 
steppe), exterminating mainly the young. Chiefly at this time in the 

*In North America, called "moose" — Sci. Ed. 



216 

steppe and desert, but also in tundra, wolves hunt pregnant ungulates 
(dzheiran, saiga, roe deer and [other] deer). At the time of birth of the 
young, wolves gather near the place of calving, where they kill both 
adults and young. 

After melting of the snow and the beginning of spring reproduction 
of animals (end of April— May) wolves transfer their feeding mainly to 
vertebrates of moderate and small size. In June, the young begin to feed 
mainly on animals, and the mother's milk is only a secondary food for 
them. In June, wolf-cubs already can walk independently to water- 
holes. From August on, attacks of domestic livestock become common 
(KJiudyakov, 1937). 

The food habits of wolves in different regions differ markedly. The 
food of tundra wolves during the snowy period is without exception wild 
and domestic [reinjdeer (mainly calves and does). They attack hares, 
arctic foxes and other animals. In the Nenetsk Nat. Okr., in the stomachs 
of 74 wolves caught in winter and spring were found remains of: rein- 
deer — ^93.1%, small rodents — 5.4%, willow grouse — 4.1%, white 
hare — 1.3%, foxes — 1.3% and fish— 6.8%. Of great importance in the 
life of wolves living at the expense of ungulates is the group attack and 
mass "cutting" of the prey into [smaller] flocks and herds. Later, wolves 
often travel to places of these "foragings" to utilize the preserved re- 
mains. Small groups of predators remain near the sea coast or by villages 
where they feed on marine flotsam along the shore, the wastes of indus- 
try, carrion, and rob the traps and snares of hunters (Makridin, 1959). 

In the summer food habits of wolves on the tundra, a considerable 
role is played by birds (in the second half — especially molting geese 
and ducks) and small rodents (lemmings and voles). [Rein]deer, espe- 
cially at the begiipning of summer (calving time), also play an important 
158 role. In the montane regions in the northeast, snow sheep, hares and 
marmots occupy a notable place in the food habits of wolves (A. 
Romanov, 1941). 

In the taiga of Karelia, wolves attack elk (young), reindeer and 
domestic livestock and eat carrion; in summer, they catch rodents, 
ground nesting birds, and sometimes lizards and frogs. They also eat 
berries, in autumn especially mountain ash (Marvin, 1951). 

In the forest of the Tatar ASSR wolves feed in the snowy period 
mainly on mammals (98% of occurrences), especially the domestic 
animals, and carrion (68%), followed by mouse-like rodents (24%) 
and hares (21%). The proportion of birds constitutes only 10% of the 
findings (as % of total number of stomachs, feces and food remains 



217 

investigated; V. Popov, 1952). In the region of the Rybinsk reservoir, 
wolves in v^inter hunt mainly elk (Kaletskaya, 1953). In the Urals, they 
feed on wild ungulates, hares, ground squirrels, domestic animals and 
birds. In winter, they eat carrion and frequently pursue foxes (Shvarts, 
Pavlinin and Danilov, 1951). In Belovezhsk Forest, they hunt chiefly 
wild ungulates (48%), especially wild boars (21%), roe deer (18%), 
[redjdeer (6%). Remains of domestic animals were encountered in 28% 
of the stomachs. They frequently catch brown hares (16%). In the 
warm period of the year, the significance of ungulates decreases to 
40% (in spring) and 31% (in autumn). At the same time, the role of 
domestic animals in their food increases from 32% to 42% (sheep 
particularly are caught). Wolves here hunt dogs relatively rarely (Gavrin 
and Donaurov, 1954). 

In the forest-steppe of the central chernozem regions, domestic 
livestock, hares and small rodents comprise the principal food of wolves. 

In steppe regions, in 56 cases (stomachs, food remnants) first place 
was occupied by mouse-like rodents (35%), then carrion ( 1 7%>— mainly 
in the winter), dogs, calves, sheep, goats, swines (16%). In the stom- 
achs, remnants of hares (8%) and ground squirrels (5%)) were often 
found; in one case, red fox. Proportion of birds (mainly domestic geese) 
constituted 4%. Occasionally wolves eat lizards and insects (dung 
beetles). Plant food are represented by berries, strawberries, pears and 
apples (windfall). In the Usmansk forest, the usual prey of wolves 
consists of deer, beavers, hares and raccoon dogs (Table 13). In sum- 
mer and autumn, remains of dogs (38%), hares (18%)) and sheep (13%) 
are found in their food (Mertts, 1953). 

Vertebrates are encountered in 92.2%) of the cases in food habits 
of Ukrainian wolves (Komeev, 1950). 
159 Foods consist of mammals — ^90.7% occurrence, birds — 12.9%), 
reptiles — 5.5%, amphibians — ^29.6%, fish — 18.5%, insects-^6.2%), 
plants — 48.1%). Among mammals, domestic animals are encountered in 
48.9%) of occurrences, and game species in 32.6%), including hares — 
22.4%, roe deer— 10.2%, mic^— 14.2%, voles-^2.8%, shrews— 6.1%. 
Among domestic animals dogs occupy 18.2%. 

Up to 10 water voles {Arvicola terrestris) and 15 common voles 
{Microtus arvalis) were found in the stomach of one wolf. An espe- 
cially great importance in the food of wolves is assumed by small 
rodents in the years of their mass reproduction (Migulin, 1938). Among 
birds, mallard {Anas platyrhyncha)*, domestic hens and geese 

*Sic: should read platyrhychos — Sci. Ed. 



218 

158 Table 13. Food habits of wolves in the Usmansk forest (112 stomachs and 63 

feces, Merits, 1953) 



^^■^^-^^.^^^ Season, data 


Winter 




Summer 




^^^"^^„^^^ 


Number of 


% 


Number of 


% 


Food type ^""-^^ 


„^ occurrences 




occurrences 




Deer 


37 


21.1 


3 


5.3 


Beavers 


27 


15.3 


3 


5.3 


Hares 


34 


19.4 


10 


17.9 


Mouse-like rodents 


14 


8.0 


5 


8.9 


Raccoon dogs 


7 


4.0 


1 


1.8 


Domestic dogs 


20 


11.4 


21 


37.5 


Sheep 


7 


4.0 


7 


12.5 


Swine 


4 


2.2 


1 


1.8 


Domestic birds 


4 


2.2 


3 


5.4 


Carrion, bait 


17 


9.6 


— 


— 


Plant remains 


5 


2,8 


-) 


3.6 



predominate, among reptiles — ^lizards and among amphibians — ^toads. 
Among fish, the pike (Esox lucius) was found, being caught by wolves 
in water-meadows (in times of flood). Other kinds of fish are caught 
along the shore. Among insects, beetles (100% of occurrences), 
Orthoptera (48%) and Hymenoptera (44%) predominated. Among the 
plant food, berries of buckthorn (Rhamnus cathartica; up to 389 in one 
stomach), black-berried nightshade {Solarium nigrum; up to 9082 seeds 
in one stomach), berries of lily of the valley (Convallaria majalis; up 
to 486 seeds in one stomach) and fruits of pear (Pints communis; 140 
seeds in one stomach) were met with (Komeev, 1950). In addition to 
animal foods, many grains of maize were found in wolves' stomachs 
in the Caucasus, and in Kiev district, the fruiting bodies of mushrooms 
of the genus Tricholoma (Komeev, 1950). 

In the Caucasus preserve the frequency of occurrence of various 
groups in the food of wolves was as follows (Teplov, 1938): mam- 
mals— 90%, ungulates— 81%, wild boar— 38%, deer— 16%, tur— 12%, 
chamois — 12%, roe deer — 7%, rodents (hares and mouse-like) — ^90%, 
carnivores (bear, fox, marten) — ^3%, birds (chiefly Caucasian black- 
cock) — 7%), fruits including berries (pear, apple sweet cherry, viburnum, 
blackberry, dog rose) — 12%. 

In the Uralo-Embensk desert, the summer feces and food remnants 
of wolves (268 examples) consisted of 37% mouse-like rodents, jer- 
boas — 9.1%, ground squirrels — 13.6%, hares — 2.6%, hedgehogs — ^2.9%, 
domestic livestock — 8.8%, birds — 15.8%, reptiles — 0.3%, insects — 1.1% 
and plant remains — 5% (Kolosov, 1935). 



219 

In the northern zone of Kazakhstan the prey of wolves in summer 
are small rodents (especially water voles), hares, young and molting 
ducks, young blackcocks and willow grouse, and rarely roe deer and 
domestic livestock (sheep). During times of drying-out of lakes, water 
birds and water voles are especially available to wolves and hence 
become the principal food, on which they also feed their litters of 
whelps. In Kazakhstan the wolves also readily eat the fallen apples and 
pears, and visit melon-fields. In winter in northern Kazakhstan, wolves 
hunt domestic livestock, roe deer and eat carrion (Afanas'ev et al., 
1953). Around Lake Kurgaldzhin, wolves live all year round in the 
reeds. In summer they feed on water voles and waterfowl, especially 
during the time of molt. In winter, they live at the expense of wild pigs, 
following them in paths which the swine tread in the snow. 

In the Betpak-Dala desert, wolves feed in summer on dzheiran, 
jaigas and hares; they also eat gerbils, jerboas, turtles and insects. In 
autumn and winter, the saigas and dzheiran which overwinter here, as 
well as the domestic livestock suffer greatly from wolves. Mammals 
predominate in wolves' food in southern Pri-Balkhash (92—100% of 
occurrences), among them ungulates (16-100%) and rodents (10-84%) 
predominate. Among ungulates, wild swine (10-50%) and roe deer (5— 
100%) are of greatest importance. The wolves frequently eat tolai hares 
and muskrats. Around water bodies, they eat fish (mainly саф). Rarely 
they attack foxes, and among birds, mainly waterfowl and pheasants 
(Table 14). 

In the lower Hi river the infrequent remains of domestic animals in 

wolves' food is explained here by the abundance of wild foods. Feces 

160 containing soil indicate the necessity to wolves of melkozem [fine 

earth] especially during the transition from summer foods to winter 

ones and vice versa (Afanas'ev et al., 1953). 

Domestic animals have important significance in the food habits of 
Turkmenian wolves; as does the wild fauna, especially ungulates which 
play a great role, particularly for the Badkhyz wolf population (south- 
em Turkmeniya). This is, obviously, connected with the richness of 
Badkhyz in wild ungulates. The particular attention of wolves here is 
concentrated on dzheiran as the largest amount of the more available 
species. Adult dzheiran are caught mainly at waterholes by wolves 
concealing themselves behind dense thickets. Cases of attacks on kulans 
by wolves are also known, especially during winter snowfalls and on 
ice-covered ground. Small animals, especially reptiles, are known to be 
significant in the food habits of wolves in the Badkhyz (Heptner, 1956). 



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221 

Wild ungulates (goats, arkhar sheep, roe deer, maral) and domestic 
livestock serve as the main food of montane wolves in Kirgiziya. Wolves 
follow them into the high mountains in summer, and in winter, they 
descend to valleys. In summer, wolves readily and successfully hunt 
marmots, concealing themselves and waylaying them; they eat also 
small rodents, birds and carrion (Kuznetsov, 1948). In the Dzhungarsk 
and Zailiisk Alatau, the marmots frequently serve as the main food of 
wolves (Afanas'ev et al., 1953). 

Change in the quantity of the chief foods or the conditions under 
which it is obtaining in different years strongly changes the character 
of wolves' food habits. In winters with much snow and particularly 
with severe and protracted snowcrust, wolves sometimes almost com- 
pletely exterminate the wild ungulates (especially roe deer), even over 
great areas. Thus, during the winter of deep snow in 1940/1941 in 
northem Kazakhstan, wolves almost exterminated roe deer in Kustanaisk, 
North-Kazakhstansk, Pavlodarsk, Kokchetavsk, Akmolinsk, and East- 
Kazakhstansk districts entirely. In the Presnogor'kovsk region of 
Kustanaisk district alone, remains of more than 300 roe deer killed by 
wolves were found in this year. In Naurzumsk preserve mass mortality 
of roe deer was noticed after the occurrence of a strong snow crust 
(16th March). This recurred in the snowy winter of 1948/1949. In the 
winter of deep snow in 1947/48 wolves attacked even adult male wild 
boars which they do not try to touch under ordinary conditions (Afanas'ev 
et al., 1953). Frozen ground and protracted continuous snow cover* in 
the steppes and deserts and the crusted snow in the forest zone show 
similar effects. In all cases, the number of ungulates shaфly decreases 
and in subsequent years its significance in the foods of wolves de- 
creases. 

In some conditions, wolves feed on completely unlikely food. Thus, 
during the mass outbreaks of locusts in the Kizlyarsk steppe in the 
161 1920's, the feces of wolves consisted entirely of the remains of these 
insects (V.G. Heptner). 

Home range. The way of life of wolves and their methods of 
utilizing territory differ in different seasons and are not the same in 
populations inhabiting different geographical zones and regions. Every- 
where, two main seasons are distinguished, when wolves lead differing 
mode of life. In the warm period of the year, in the time of parturition 
and rearing the cubs, adult wolves live in pairs (families), while 
pereyarki (young of the previous year) [yearlings] live singly or in 

*In Russian, dzhvt — Sci. Ed. 



222 

small groups, usually not far from the parents. Both groups are fully 
established or accomplish only short migrations mainly at the end of the 
season. 

By the cold time of year, the yearlings join the adults with this 
year's young. Thus a "pack" of wolves is formed, representing, in fact, 
a family group consisting of the parents, juveniles (this year's litter) 
and yearlings which have not yet reached sexual maturity. It more 
rarely happens that older animals, usually males 3-5-year old, which 
have been left alone for one or other reason, join such a "pack". 
Strange animals, bom to other parents, are not admitted into the pack 
and are regarded as enemies. 

The average size of a pack is 1—2 adults, 3-6 juveniles, and 1—3 
yearlings, i.e., 5—1 1 beasts. The largest pack known was composed of 
16 beasts (2 adults, 10 juveniles and 4 yearlings). On the European 
tundras, packs are usually composed of only the parents and young — 
mostly 5-7 animals (maximum 10 wolves). Apparently, on the tundra, 
yearlings do not always join their parents (Makridin, 1959). In 
Arkhangel'sk district, the pack is usually composed of 7-8 and rarely 
of 10-12 animals (Semenov, 1959). In wolf packs in eastern Siberia, 
there are usually 7-10 animals (Khudyakov, 1937; Timofeev, 1949). In 
Ryazansk district, in a total of 14 years of observations only once was 
a pack of 1 1 encountered (1946), and 2 times of 10 wolves (1945 and 
1947; Kozlov, 1955). In Tatariya, over several decades of observation, 
no pack was found consisting of more than 12 wolves (V.A. Popov). 
In Belovezhsk Forest, where wolves were numerous, a pack of 12 
wolves was found in 1946. Apparently, in this region, packs of 15 and 
16 wolves may be present, but more often, they consist of 8-10 animals 
(Serzhanin, 1955). In Kazakhstan, 3-12 wolves occur in the pack, and 
rarely — 15. In the lower Hi river, packs of up to 10 wolves are met with 
(Afansa'ev et al., 1953) (Table 15). 

162 Table 15. Number of wolves in a pack in the lower reaches of lii river 

Number of wolves "o 



Time of \. 
observation \^ 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10* H S. 


> N 
< -^ 


October— December 

January 

February 


8 
14 

2 


4 
6 
3 


6 

1 

3 


3 


3 


— 


2 


1 


— 


28 28 

21 21 

8 8 


3.3 
1.4 
2.1 



*Figures in this column are clearly in error; they should be (from top down) 1, — , — . 
— Sci. Ed. 



223 




162 Fig. 55. Hunting areas of wolf pup litters in winter in Ryazan district in the 
region of Oka preserve (from Kozlov, 1952). 
1 — ^location of litter, 2 — ^hunting area of litters, 3 — consistent passage of wolves, 
4 — preserve boundaries, 5 — ^migrations of wolves for prey in summer. 

The average number of cubs in a pack in Aktyubinsk and North- 
Kazakhstansk districts in December— January was 6.3 animals (of seven 
packs; Afans'ev et al., 1953). 



224 

162 The pack is sometimes divided into groups. Thus, in the winter of 
1944/45 a pack of 10 wolves, living in the region of Oka preserve, was 
often divided into two groups, each of 5 wolves, hunting independently 
at a distance of 8-10 km from one another. One group was led by the 

163 mother and the other by the father (Kozlov, 1955). The winter pack 
completely dissolves only with the deaths of both parents. 



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Fig. 56. Distribution of wolf pup litters in the region of Oka preserve (Ryazan 

district; after Kozlov, 1952). 

1 — place and year of litter, 2 — preserve boundaries. 



225 



On the tundra, wolves feeding on prey from the traps of hunters, 
carrion and bait at villages or edible flotsam by the sea, do not live in 
packs, but singly or in pairs. Packs apparently cannot be sustained on 
such food (Makridin, 1959). Old wolves incapable of reproduction live 
also alone in winter (Kozlov, 1955). Sometimes, killing of weak or 
diseased animals is observed in the wolf pack. Thus, wolves usually 
tear apart and eat an animal which eats a strychnine bait and goes into 
convulsions (Shvarts, Pavlinin and Danilov, 1951). 

As a rule, packs live separately and on meeting, they relate to one 
another with hostility, although cases of serious clashes, fights, or 
especially "battles" are unknown. Wolves which enter into the hunting 
territory of a strange pack of wolves leave it at once upon their first 
meeting with the owners of the territory, or upon discovering their 
"scent marks" (see below). In rare cases, separate packs temporarily 
164 unite and together attack a herd of wild and domestic ungulates; in this 
case, the size of the united group may reach 20 and more animals. Such 
a union is always transitory and of short duration, and usually happens 
in places of crowding of ungulates — near large migrating herds, in the 
places of mass calving, and so on. In the Mongolian Republic, such 
joining of wolf packs was observed around large herds of dzeren 







hkML^ lb; V'i ' i' "i\ 



ft > 



-^ 





Fig. 57. Birch clear-cut in steppe — denning place of wolves. Krasnoyarsk 
territory. May 1959. Photograph by V.V. Kozlov. 



226 

[Mongolian gazelle, Procapra gutterosa] (Bannikov, 1954). During 
periods of pack life, wolves without growing young are very active and 
in some places perform extensive wandering, following the migrating 
wild ungulates or herded domestic livestock. 

In the second half of winter and to its end, the mothers with their 
by this time maturing yearlings separate from the pack and wander in 
pairs or groups. The young wolves with the still not mature yearlings 
in the lead, remain together, and in their absence, they disperse into 
groups of 2—3 animals or wander singly. 

The summer sedentary life of wolves begins, in the middle zone, 
usually in April, with the arrival of pregnant females at den sites. 
Whelping places (dens) are permanent; they are only changed in case 
of extreme necessity, particularly alterations in the surrounding land- 
scape unfavorable to wolves. The den is situated either at the center or 
in one of the parts of the hunting territory of the family (see below). 
Pregnant female wolves usually return to their old dens, while the 
young that are pregnant for the first time, search for new lairs which 
are, as a rule, not far from the place of their birth. At that particular 
time the winter packs of wolves finally break up and the juveniles, 
having now become yearlings, separate from the parents and live inde- 
pendently, usually not far from the mother's den. 

Different wolf families have mutually exclusive hunting territories, 
and the simultaneous living of two families in one territory is, as a rule, 
impossible. Under certain conditions, at least in winter, this rule is, 
however, not followed. Thus, in winter of 1920/1921, at the time of 
great food scarcity (dzhut), a great number of horses wintered on 
pasturage in glades in steppe areas and large meadows among the 
165 clear-cuts of the Ara-Karagai forest (b. [former] Kustanaisk, Turgaisk 
governance). The wolves, numbers of which were especially high in 
that year, gathered thither from a large region and killed the horses. 
The number and density of wolves in Ara-Karagai was so great that 
there was no separation of hunting territories of the packs and perma- 
nent packs were not really present (V.G. Heptner). 

The territory of the pack and its boundaries are distinguished by 
special characters — scent marks— places where wolves urinate. They 
are usually situated near things — crocks, boulders, a single tree, the 
skull or skeleton of a large animal — that are easily distinguishable from 
afar. On passing by such a place, wolves must visit them and leave 
their own marks, indicating that the spot is occupied. These "urine 
points" are the best places for setting traps. 



227 




Fig. 58. Small cliff in the forest-steppe under which was a den with 6 wolf pups. 
May 1958, town of Khakassiya, Krasnoyarsk territory. Photograph by 

V.V. Kozlov. 

The size of the hunting territory of the wolf family, and conse- 
quently the distances between summer dens, depends on the availability 
of food in the territory (stocks of summer food of wolves), cover, and 
many other conditions. In the middle zone, dens are located usually no 
closer than 6—12 km from each other (radius of the hunting territory is 
3-6 km; Kozlov, 1955). In the Ukraine, summer hunting territories of 
wolves are smaller. Dens here are situated near one another, sometimes 
in neighboring ravines or in one woodlot (Komeev, 1950). Exception- 
ally, two female wolves sometimes give birth to their young in a single 
den. Thus, in the Sinyavsk region of Sumsk district in 1948, 8 wolf 
pups and 2 old female wolves were found in one den. Among the cubs, 
four were noticeably larger and obviously belonged to another litter 
than the other four (Komeev, 1950). 

In the taiga of eastern Siberia, the average radius of the hunting 
territor>' is approximately equal to 10 km in summer (Khudyakov, 
1937), while in steppe and especially in desert zones it is larger and 
the wolves' dens are located no closer than 15—20 km from one an- 
other. Only a few exceptions are known. Thus, in the Vishnevsk region 
of Karagandinsk district, three burrows of wolves were found at a 
distance of only 500 m from each other. Two pups were found in one 



228 

burrow, 5 in another and in the third, 6 pups with eyes open were found 
(Afanas'ev and others, 1953). In the middle zone, cases are known 
(very rare) when two litters lived at a distance of only 2 km from each 
other. These were 2 female wolves with their young, the males of which 
166 (both) died in the spring (Sokolov, 1951). At the same time, there is 
some indication that when neighboring dens are close, the female wolves 
ate each other's young (Danilov, 1945). In the mountains, dens are, as 
a rule, located nearer to each other than in forest steppe and desert 
zones. 

At the time when the juvenile wolves are present in the den, the 
parents usually do not hunt in the immediate neighborhood, but go out 
to the farthest parts of their hunting region. Thus, in 1942 in the Altai, 
in snow which had fallen on May 20th it could be traced that parents 
went a distance of 12—25 km from the den for their prey, at a time when 
wild ungulates and grazing livestock were living in their neighborhood 



-if'' T 



Fig. 59. Tracks of two wolves, passing a day lair, tlie place of their daily rest. 
Khakassiya, Krasnoyarsk territory. 12 December 1956. Photograph 

by V.V. Kozlov. 



229 

(Nasimovich, 1949; Afanas'ev et al., 1953). In the middle zone, the 
radius of summer hunting chases does not exceed 5—10 km (usually 5— 
6) (Sokolov, 1951). Thus, in Ryazansk district, in the period when the 
young are being fed, the aduh wolves hunt usually no closer than 5— 
7 km from the den and do not touch livestock grazing within the limits 
of this zone. In the summer of 1939 in the Charussk forestry [area], a 
den of wolves with a litter was found 1.5 km from Chemaya Rechka 
village; livestock from this settlement were pastured in the immediate 
proximity of the place of the litter, but no losses caused by wolves were 
noticed. At the same time, complaints about wolves were constantly 
being lodged from those who were raising livestock at a distance of 7 
km from the litter's location. After destruction of the litter, there was 
not even one case of attacks by wolves on livestock. Only yearlings 
sometimes break the "rule" of not touching livestock near the den of 
the wolf family (Kozlov, 1955). 

In a convenient den, located in a favorable place, the litter lives the 
whole time until autumn and the beginning of the nomadic period. 
However, if the den is established by the adult wolves far from a water 
source (2—5 km away), it is used only until the young are weaned from 
the mother's milk. Later, the litter resettles nearer water. The particu- 
larly great need for water is manifested in the young at their transition 
to a meat diet. Upon the drying up of a water source, wolves also lead 
or transfer the young to a new den. During a period of drought in 
Kazakhstan, wolf pup litters are concentrated near the few intact water 
sources, devastating their neighborhood, while some litters perish due 
to starvation and especially the deficiency or absence of water (Afanas'ev 
et al., 1953). 

A den-burrow built in open, non-overgrown ravines is also changed. 
It is only used so long as the wolf pups do not come out of the burrow. 
Later, the parents lead the litter to more hidden places (thickets of 
shrubs and weeds, forest islands and sometimes to winter crop fields 
where bread grain is grown)*. In the Oka floodplain, the adult wolves 
frequently led the growing litters to places with an abundance of small 
fruits — raspberries, strawberries, red billberry [Vaccinium vitis-ideae] 
and others (Kozlov, 1955). 
167 The she-wolf sometimes builds a temporary nest for whelping, 

particularly when the places of the main nests are flooded by melt- 
water, or when the she-wolf is frightened in some way before parturition 

*Probably refers to winter wheat — Sci. Ed. 



230 

from the main nest. The duration of their stay in the temporary shelters 
is determined by the local conditions, but before the young of the 
current year become independent of their mothers, the litter usually 
changes several temporary dens. In these cases, the she- wolf accompa- 
nies the young to the main lair when they are already quite independent 
(Sokolov, 1951). 

If wolves in a den were disturbed, particularly if their litter was 
touched, they usually abandon the den, carrying the surviving young to 
a new place. With closing of the entrance to the den (burrow), the 
parents usually dig it out at night and carry off the pups. At such times, 
neither the female or male wolf defend the nest and do not attack its 
invader, even if the nest is destroyed and the pups taken (Kozlov, 
1955). While the adults are busy raising the litter, the yearlings remain 
in the region of the litter, but they do not approach the den closely. 
They live alone, or in groups of 2-3 animals and hunt independently. 



-'^m 



V%- '4V 






Fig. 60. Resting place of wolf in forest-tundra. Photograph by V.P. Makridin. 



231 

At the start of raising the litter, the old wolves remain extremely 
secretive; they usually do not howl in the region of the den until July. 
In June in the south, and in July in the north, the demands of the young 
for food noticeably increase, and the parents significantly expand their 
hunting territory. Precisely at this time, the young begin to give voice, 
apparently because above all they often feel hungry. After the young 
develop, their elders begin to howl upon their return from the hunt, and 
the juveniles answer the old ones. At the same time, cases of attacks 
by wolves on livestock become more frequent in the vicinity of the den. 
Thus, in 1938 in Kalininsk district, a horse was killed only 500 m from 
168 a den, and the she-wolf for several days led the litter to the carcass to 
feed. Near Bologi, a maturing litter of wolves assiduously tore up sheep 
and calves, not farther than 2 km from their den (Sokolov, 1951). 
Beginning with this period, together with the young (juveniles) may be 
found the yearlings which join the family till the time of the next rut. 
In July (in the south) and August-September (in the north) the maturing 
young wolves attain half the size of the adults and begin to participate 
in the hunt with their parents. 

The size of the hunting territory continues to increase in autumn, 
and the wolves gradually convert to a wandering life. Henceforth, the 
old wolves stop bringing food to the den; on killing a large prey, the 
parents lead the young to it, and after satiation, the family returns to 
shelter by the morning (Khudyakov, 1937). 

When the milk teeth of the young are replaced by permanent ones, 
the wolf family begins to hunt together. At first, however, wandering 
is restricted to the hunting territory. Roaming increases with the fall of 
snow and the termination of livestock pasturing. In the middle zone, the 
family rarely returns to the den site in the winter, establishing instead 
diumal lairs (Kozlov, 1955). 

In forest regions of Arkhangel'sk district, fi^om September wolves 
already lead a nomadic way of life, making long movements and spend the 
day where they are met by the dawn. At this time, each wolf family 
occupies a definite hunting region, leaving it only in case of prolonged and 
shaф hunger or continuous pursuit by humans. Tundra wolves are, at this 
time, still more active. In eastern Siberia in winter, wolves wander widely, 
following wild ungulates, mainly roe deer. Beginning in November, they 
hunt ungulates almost exclusively; making chases or driving the roe deer 
onto the ice, capturing them in their beds; and frequently bringing izyiibr 
[Manchurian wapiti] to "bay". These predators do not remain for long in 
one place; the hunting territory of the pack sometimes reaches a diameter 
of 70 km (Khudyakov, 1937; Timofeev, 1949). 



232 



Ьб^ 




'* '^I^Ш^^'^ 





Fig. 61. Tracks of a pair of desert wolves, hunting a saiga. Central Betpak- 
Dala, Kazakhstan. 7 January 1957. Photograph by A. A. Sludskii. 

169 In the central districts, during snowy periods of the year, the hunt- 
ing region of a wolf family is also expanded. In Ryazansk district, the 
family hunting range of wolves had a diameter of 25 to 40 km. In forest 
and forest-steppe districts of the European part of the USSR, wolves in 
winter perform movements of about 15—30 km during the night and 
when hungry, up to 50-60 km (Kozlov, 1955). In Voronezh and con- 
tiguous districts, wolves leave the permanent summer habitats in 
November, when the radius of their hunting activity does not exceed 5 
km and begin to wander widely, dividing up into small groups, carrying 
out invasions of livestock yards. At this time the radius of their hunting 
range increases to 15 km and more (Mertts, 1953). In forest-steppe 
regions, the greater part of the animals concentrate in forests, where in 
winter the food is more abundant, and they usually occupy permanent 
hunting ranges. With the coming of spring, they again wander back to 
the fields (Mertts, 1953; Barabash-Nikiforov, 1957). 

Pack formation in the snowy time of year makes it easier for 
wolves to obtain food, especially the young (see below). In places with 
deep porous snow cover, the pack moves in a line, always with an adult 
wolf in front. In places with little or no snow, for example in the lower 



233 




Fig. 62. Tracks of a desert wolf following a saiga. Betpak-Dala desert, 
Kazakhstan. 19 December 1956. Photograph by A. A. Sludskii. 

Hi river, the pack frequently moves in a helter-skelter fashion (Afanas'ev 
et al., 1953). 

In searching for prey, wolves move around their hunting range 
along a definite route (Fig. 55), using one and the same trails for 
extended periods, restoring them after snowfalls; during this, the wolves 
without fail find their old tracks, and follow them exactly, track by 
track. Trails are laid out along the banks of rivers and shores of lakes, 
in places of passage between plantations, ravines overgrown with shrubs, 
etc. Widely used are roads and paths made by humans (Semenov, 
1954). 

Wolves on the tundra lead a pack way of life for the greater part 
of the year, because attacks on [reinjdeer herds and the pursuit of 
stragglers from the herds are more successful when hunting in a group. 



234 

In the taiga zone also, particularly in the second half of winter, wolves 
do not often find prey sufficient for feeding the whole pack, and thus, 
cooperative hunting becomes temporarily unfavorable, and the wolves 
hunt in pairs or singly, gathering in the pack only for diurnal rest. 
Sometimes the pack is broken up and for a week or two the wolves live 
in collections of not more than 2-3 animals (Semenov, 1954). 

With the beginning of the rut, the mature wolves drive away from 
themselves the young and yearlings that up till now have lived with 
them. After heat which continues for 5-10 days, and the end of rut, the 
wolf family gathers together again for a short time for cooperative 
hunting of large prey. In the north, approximately in April, such packs 
170 gather on the main trails along which the [rein]deer herds move to their 
summer pastures, and in following these herds, they move to the tundra 
and to places of littering of the young. 

Burrows and shelters. Different protected places serve as diurnal 
lairs (resting places) for wolves. In cold, windy and damp weather, the 
wolves prefer the more protected places, while in dry, calm and warm 
weather, they willingly lie down in the open. In autumn— winter-spring 
period, when the wolves lead a nomadic, or in general a more active 
mode of life, they lie down wherever they are at daybreak. In summer 
time, they are associated with dens which they must build for the young 
(see above). 

In constructing the den, the she-wolf usually makes use of natural 
shelters — ^fissures in rocks, cliffs with overhangs by the banks of rivers 
and slopes of ravines, holes thickly covered with plants, thickets of 
saplings or bushes, "brushwood", or fallen branches, weeds, etc. In 
many cases the nest is constructed in the burrows of various animals — 
foxes, badgers (in the forest zone), marmots (in steppes, montane 
meadows), arctic foxes (in tundra), etc. Wolves usually widen the 
adopted burrow and partly remake them. In rare cases, the she-wolf 
digs a burrow, which is usually shallow and short, with 1—3 openings. 
The burrows are built on hillsides with soft, mainly sandy and sandy 
loam soil. In the majority of cases, the carnivore's nests are situated in 
dry sections. 

The den, as a rule, is constmcted not more than 500 m from a 
water source (lake, river, creek, spring, deep hole with water, pond, 
etc.); to it one-two trails lead, becoming especially noticeable when the 
young begin to leave the burrow. The approach to the water source is 
often concealed. Near the trails around the den, a large number of 
resting places of wolves, places of play for the cubs, and food remains 



235 

are encountered. From the den there usually emanates a shaф odor of 
urine and of rotting food remains. The food remains attract such birds 
as magpies, ravens and others to the nest. By their cries, it is possible 
to locate with precision the nests of wolves with litters. 

Convenient places for dens are few, and from year to year they are 
occupied usually by one and the same pairs of wolves. If the male dies, 
he is replaced by a yearling in the pair, and if it is female — by a young 
she-wolf. Empty dens are therefore rare, and they are only left due to 
the fundamental changes in the condhions of the surroundings (felling 
of the forest, fire, marsh drainage, etc.). Thus, in the Charussk forestry 
[area] of Ryazansk district, wolves established a den for four succes- 
sive years (1936-1939) in one and the same woodlot, although every 
autumn a drive was mounted against the litters appearing there, when 
a portion of the young were killed, and in 1937 the mature male was 
also killed (Kozlov, 1955). 

The wolves more frequently select for dens, places rarely glanced 
at by humans, but they construct them near domiciles, paved roads, 
railways. In 1941 a wolfs nest with young was found within 200 m of 
the Moscow-Leningrad highway (Sokolov, 1951). In Zheltukhinsk re- 
gion of Ryazansk district, a she-wolf built a den in part of a field 
overgrown with weed only 1.5 km from the village. Another wolf den 
was discovered 10 m from the road-bed on the Moscow-Ryazan rail- 
way, under a pile of snow-screens (Kozlov, 1955). A she-wolf with a 
litter lived in the middle of a peat-bog not far from a corduroy road 
along which carts frequently passed (Manteifef, 1949). In Ryazansk 
district a den was found in a small but dense birch clear-cut not far 
from a large village (regional center of Erakhtur) and in immediate 
proximity to a large road with much automobile traffic (V.G. Heptner). 
In the depth of large forest massifs, wolves do not usually make dens. 
In Tatariya, all lairs were situated not further than 500 m from the 
forest edge, usually along ravines where there is water (V.A. Popov). 
171 In different landscapes, the situations and construction of dens are 

somewhat different. Tundra wolves use shelters of three types: a) land 
burrows, which they build themselves or adapted from ones of foxes, 
and rarely, arctic foxes. The burrow consists of an entrance trench 1.5 
m in length, an underground corridor 50-60 cm wide, and length 2 to 
10 m, and a nest chamber 150 x 100 x 70 cm; the chamber is situated 
at a depth of 1.5 to 3.5 m, and usually contains no nest bedding; b) 
burrows under rocks of nearly the same construction, but usually shorter; 
c) different natural shelters (caves, fissures in rocks, etc.). The wolf 



236 



't'*-».!'-? 



Fig. 63. Tracks of a wolf and a she-wolf Uzhursk region, Krasnoyarsk territory. 
December 1957. Photograph by V.V. Kozlov. 

den is usually situated on the southern and south-eastern slopes of river 
banks or hills; they are better warmed by sun, and the snow thaws more 
quickly there (Semenov, 1954). 

On the Kaninsk tundra, wolves are mainly confined to hills in summer. 
In Lena-Khatanga territory, burrows are usually no longer than 1.5 m and 
not deeper than 1 m (A. Romanov, 1941). In Anadyrsk territory, wolves 
give birth to pups mainly in land burrows (Sokol'nikov, 1927). 

In the northern forest districts, wolves settle on dry ridges in the 
middle of marshes, on islands of coniferous forest with abundant fallen 
branches in coniferous saplings. They prefer burned-over mossy bogs, 
especially in the difficult-to-penetrate margins overgrown with impass- 



237 

able small trees. Wolves settle in wet areas covered with alder woods 
and on islands of dry, but dense, forests with spruce understory, which 
are rarely visited by humans (Semenov, 1954). 

Wolves living in the forests of Ural and Siberia, usually have dens 
for their litters situated directly on the surface of the earth, under 
bushes, overthrown trees, etc. With wolves of the open steppes and 
forest-steppes, the denning place is more often built in a burrow, as a 
rule in a foreign one — usually of fox, badger, or marmot (Khudyakov, 
1937; Shvarts, Pavlinin and Danilov, 1951; Laptev, 1952). In the steppes, 
burrows were also found distant from water sources — up to 12 km 
(Borzinsk steppe). They were constructed in open, usually elevated, 
places. In southeastern Siberia, a wolf den was found on a promontory 
by some falls where only a few birches grew, but, it was dry there and 
the she-wolf had a good field of vision, which protected it from зифпзе 
attacks (Khudyakov, 1937). 

In Belorussia, a litter of 6 pups was found in a fallen hollow alder 
tree on a small hillock in a thicket of small alder trees (Serzhanin, 
1955). 

In the central chernozem regions, dens of wolves are situated in 
young and old broad-leafed and mixed forest islands, in steppe ravines 
and gorges overgrown with shrubs, and in reeds and willow thickets in 
172 marshes, along the margins of bogs, and sometimes even in old straw 
stacks and in the remains of corn-shocks. In the steppe section of 
Voronezh district, the wolves raise their young mainly in "yarugas" — 
ravines densely overgrown with small oaks. In forests, dens are 
constructed under uprooted wind-fallen trees, sometimes in burrows of 
foxes and badgers. In fields, the predators use old military trenches, 
and in ravines — ^in holes and cavities. 

In time past, there were repeated cases of pups being transferred 
by she-wolves to tall cereal grains and construction here of dens in the 
immediate proximity of villages (Merits, 1953). In the Kantemirovsk 
region of Kamensk district, in the autumn of 1953, was found a burrow 
with two exits situated in a rye field where the wolves successfully 
brought the young. Here such dens are not rare (Barabash-Nikiforov, 
1957). In Chemigovsk district, a she-wolf built a den and brought the 
pups to the hemp plot of a personal holding of a collective farmer 10 m 
from his house. In the Kovertsovsk region of Volynsk district, a litter 
of wolves lived in a rye field, 300 m from the cottage. The den was 
constructed in a hole by the side of an uprooted stump. Such cases are 
known in Sumsk, Zhitomirsk and other districts. In Donetsk district, a 



238 

wolf den was found 150-200 m from a very busy paved road (Komeev, 
1950). 

In the forest regions of the Ukraine, dens are frequently built on 
islands among sphagnum and other bogs, in dense undergrowth of 
saplings and in forest islands with dense shrubbery among fields 
(Komeev, 1950). In forest-steppe regions, dens are situated in deep 
forest ravines, in sparse forests along river valleys and in impassable 
alder wood along tussock bogs (these places are called "volchatniks" 
[belonging to wolves]). 

In forest-steppe zone of Omsk district, the wolf selects for dens 
ridges among marshes, ravines overgrown with bushes, thickets of sage- 
brush in virgin land, and forest islands (Laptev, 1958). In the steppe 
zone of the Ukraine, dens are more frequently present in deserted 
gorges with thickets of spiny bushes and tall weeds in depressions 
between sandy hillocks, overgrown by willow and other bushes, while 
on the floodplain of the Dnepr and Dnestr- — ^along thick stands of reeds 
and other dense places (Komeev, 1950). In the forest-steppe and steppes 
of Kazakhstan, den burrows are found in the forest, at the edge, and 
in open places, usually among tall weeds or bushes at a distance less 
than 1 km from a water source. In the forest preserve "Borovoe", a 
wolf den was situated in a cave on a hill covered by a pine forest; this 
den was thrice destroyed and near it, she-wolves were twice killed. 

Dens dug by wolves in the steppe are usually 2—5 m long, rarely 
more, with one or two exits, and the nest lies at a depth of 1—1.5 m 
from the surface. Old, repeatedly used dens have very wide passages 
through which humans can easily crawl, and several nest chambers; 
entrances to the old chambers are frequently filled in when wolves 
prepare a new one. Lairs in the open are established under cover of 
bushes or tall weeds. In North Kazakhstan district an open lair was 
found in the reeds of an almost dry lake; the nest was situated in a large 
opening among meadowsweet [Filipendula], around which shallow water 
was present. Within such a lake, a she-wolf brought a large heap of 
cattail and reed and gave birth to 7 cubs on it. 

In the desert zone of Kazakhstan, wolves give birth to cubs both 
in burrows and in open-air dens. Burrows in the southem Pri-Balkhash' 
were found in solonchaks [salt flats]. They were 7 and 8 m long and 
1—1.5 m deep. In sandy mounds covered with saxauls, a den was 
excavated at the top of a mound under the roots of saxaul and consisted 
of a simple niche, in which 6 cubs were lying. One burrow was situated 
in a dense growth of saxauls in the Hi river valley. Here open-air lairs 



239 

were also found in reed growths and under willow bushes (Afanas'ev 
et al., 1953). In Kazakhstan, of 24 wolf dens, 14 were situated in 
burrows dug by she-wolves, one was in a badger burrow, another one 
in a cave, another in a niche and 7 were in the open (in tall weeds, rose 
willows or reeds). 
173 In Turkmeniya, wolves inhabit the plains, old stream channels in 

cliffs of the chink along the Uzboi, in ravines of the Kopet-Dag [moun- 
tains], near creeks and springs, chiefly in the shady places, under the 
overhanging rocks, etc. Sometimes, dens are situated in dense growths 
of tamarisks {Tamarix) or other bushes, and often in tugai (riparian 
vegetation) (V.G. Heptner). In Armenia, these predators usually use 
narrow caves, fissures and ravines for dens (Dal', 1959). 

In the montane steppes of Kirgiziya, on high-alpine watersheds and 
in the foothills, wolves usually construct the den in a burrow dug in a 
ravine, under a bush, or in another such protected place. On the slopes 
of mountains, they use fissures in the rocks, overhanging shelves of 
stone, marmot burrows, etc. (Kuznetsov, 1948). 








Fig. 64. Tracks of a desert wolf in mud near a waterhole in the Gyaz'-Gyadyk 

mountains. Badkhyz preserve, southern Turkmeniya. February 1965. Photograph 

by V.G. Heptner. 



240 

Daily activity and behavior. Wolves are nocturnal predators. In 
the majority of places they lead a nocturnal mode of life in summer as 
well as in winter. 

In winter, wolf litters begin hunting in the early evening twilight; they 
hunt the entire night, sometimes traveling tens of kilometers, but neverthe- 
less often they lie down to their diumal rest hungry. In sparsely settled 
regions, hunting during the day is also observed, usually for wild ungulates 
(Komeev, 1950). In those places where wolves are active only at night, 
individual wanderers also hunt during the day. Thus, in January 1951, in 
the region of the Oka preserve, one yearling she- wolf picked out among all 
prey particularly those that were active by day (V.G. Heptner). Attacks of 
wolves on livestock were common, and especially on single horses during 
the daytime, often at the village itself, in the period of their very high 
numbers at the beginning of the 1920's in former Kustanaisk district of 
Turgaisk govemance (V.G. Heptner). With the arrival of winter, when 
lying down rather than a bed the wolf tums around until there is no longer 
any packing of the snow, in order to form a good pit-lair. While sleeping, 
wolves space themselves in a semicircle, covering their "scenter" (end of 




/' '^'Al'S^'.e 





174 Fig. 65. A wolf calmly walking in deep snow. Krasnoyarsk territory. January 
1958. Photograph by V.V. Kozlov. 



241 

the nose) with their tail. Sleep is very light, the ears are constantly either 
pricked up or drooping. Wolves sleep especially lightly in times of frost. 

In eastern Siberia, in summer, the wolves mainly stick to a noctur- 
nal mode of life, although whelps are frequently active by day also. In 
autumn, after the transition of the whole litter to a nomadic life, they 
hunt mainly at night. 
174 In southern Turkmeniya (Badkhyz), wolves lie by day in tamarisk- 
like or other shrubby thickets, and sometimes almost at the outskirts of 
settled places, always relatively close to water sources, but in sandy 
areas, at a distance of 15 km from wells (Heptner, 1956). In settled 
regions of Kirgiziya where wolves are frequently disturbed, they mainly 
lead a nocturnal mode of life; but in remote places, one may see them 
hunting during the day (Kuznetsov, 1948). Mongolian wolves hunt at 
any time of the day or night (Bannikov, 1954). 

Hungry wolves become aggressive. A huge number of cases are 
known of severe attacks of wolves on livestock. Thus, on 1 February 
1953, two wolves attacked the calf-pen of a collective farm in the 
Dryazginsk region of Lipetsk district. On hearing the noise, the people 
came running; however, their appearance did not cause the wolves to 
run away. A struggle began which ended only after one wolf was struck 
with pitchforks. In autumn, wolf litters commit audacious onslaughts on 
herds of sheep and flocks of geese, sometimes seizing their prey almost 
from the hands of the shepherds. Under ordinary conditions, wolves 
fear humans and are very cautious. They can well differentiate between 
dangerous hunter and unarmed herdsmen, in front of the eyes of which 
they frequently attack the sheep. 

Pursued wolves sometimes seek shelter in villages. In 1950, a wolf, 
having been shot from an airplane near the village of Vyselka 
Rozhdestvensko — ^the Khavsk region of Voronezh district, fled to a 
village and took shelter in the hayloft of a house near which a calf was 
lying. The wolf was discovered only after some time (Barabash- 
Nikiforov, 1957). In Kirgiziya, the wolves in winter approach villages 
and frontier posts even during the day (Kuznetsov, 1948). 

Wolves hunt in various ways: from concealment, pursuit of prey, 
and organized as a drive in groups. In summer they hunt mainly singly, 
the wolves frequently ambushing their prey, rarely spotting and pursu- 
ing prey. When attacking a herd, even individually, wolves try to frighten 
the animals, creating panic and dispersing the herd or separating one 
or several animals from it. 



242 

175 Upon attacking a herd, wolves are able to distract the attention of 
the shepherd and often conduct a very well-organized combined hunt of 
ungulates and hares, in which the functions of each member of the pack 
are not only different, but also coordinated. Breaking off from one 
another, they envelop the pursued animal from different sides and at 
last surround it. Frequently, some of the wolves pursue an animal and 
another group tries to intercept or set up an ambush in its path (Timofeev, 
1949). Once on the Malyi Cheremchan river in Tatariya, there was 
observed about 9 in the morning a unique hunt by wolves for geese: a 
wolf drove the swimming geese to a bridge, at which a hidden she-wolf 
intercepted them (V.A. Popov). 

Along black [snowless] paths or in light snow, wolves can pursue 
prey at a speed of 55-60 km per hour; with this they quickly catch even 
the brown hare [Lepiis europaeus] which runs at a speed of 40-45 km 
per hour. Wolves obtain roe deer by driving them in persistent pursuit. 
They destroy many marals by chasing them in winter into [soft] sedi- 
ment surrounding a rock, the predators patiently wait until the legs of 
the maral stiffen and it becomes unable to remain by the stones. Four- 
five wolves do not fear to attack an elk on frozen snow-crust, but with 
little snow, the adult elk is for them a dangerous enemy (see above). 
The pack can easily cope with young pigs and sows, but they usually 
fear to attack boar (Timofeev, 1949). Wolves pursue and frequently kill 
foxes but they rarely eat them. Hence, where there are many wolves, 
foxes usually become few. In the European part of the [Soviet] Union 
wolves in winter very successfully lure dogs from villages and catch 
them by feigning a cowardly flight (Komeev, 1950). Wolves at night, 
and sometimes during the day, enter villages and settlements. Thus, in 
August 1954, wolves carried away a sheep which spent the night in the 
street from the outskirts of the village of Kopanovo (Ryazansk district), 
and injured another (V.G. Heptner). Wolves follow the barking of the 
hound dogs for many kilometers and often catch them during the hunt- 
ing season. In the agricultural central-chernozem [black earth] regions, 
wolves have adapted to "mousing" behind the tractor ploughs, remain- 
ing some distance from it and catching the voles flushed out by the 
plough (Mertts, 1953). 

In searching for prey, the wolves depend mainly on their sight and 
hearing; scent is relatively weakly developed in wolves, and therefore, 
they rarely discover hidden hares or birds. But, when they come across 
a fresh track, they can easily follow it without becoming confused 
(Timofeev, 1949). Hearing of wolves is very зЬаф; the organ of hear- 



243 

ing is constantly exerted, the wolf hears and constantly registers the fall 
176 of autumn leaves from the trees. It nearly never happens that a sleeping 
wolf can be approached, while it is not very difficult to approach a 
sleeping fox. Only a replete sleeping wolf can sometimes be approached 
to within 10—30 steps. 

The sense of smell of the wolf is weaker than that of many hunting 
dogs. The wolf can scent carrion on the wind for not farther than 2— 
3 km. Because of the comparative weakness of their sense of smell, 
wolves can be taken in traps relatively easily. The sight of the wolf is 
not so complete as that of dogs. In darkness, the wolf sees better than 
the other species of the dog family because, through their way of life, 
they are by preference nocturnal animals (Komeev, 1950). Even at 



5> => 















175 Fig. 66. Diagram of the tracks of a wolf 
pack walking, as usual, one behind the other; 
when turning they separated but again went 
one behind the other. The tracks during the 
turn represent the only reliable possibility to 
establish the number of wolves in the pack — 
4 in the present case (from Kozlov, 1955). 



4S^^ 


e» "it' 




• 


v^ 


^ 



244 



Щи 




Fig. 67. А wolf on guard. Khakassiya, Krasnoyarsk territory. Winter, 1957. 
Photograph by V.V. Kozlov. 

night the wolf can easily detect the flaws in disguised traps. At the 
same time, wolves are far-sighted and they can distinguish between the 
armed hunter and the simple passerby at great distances. Upon concen- 
trating their attention on any item or occurrence, the wolf considerably 
blunts its perception of others. Numerous occasions are known when, 
fleeing from beaters, wolves came very near to shooters, even those not 
hidden. A wolf may approach with the wind, a person standing motion- 
less especially when dressed in camouflaged clothing up to 5—10 m, not 
smelling an odor or noticing a motion (Kozlov, 1955). 

On the whole, the level of nervous activity of wolves, and their 
ability to consider situations is quite sufficiently high. The behavior of 
wolves is more complex and "wiser" than is assumed by the majority 
of zoologists and hunters. When going to their dens, adult wolves 
skillfully select hidden approaches. At that time, they make numerous 
"dodges" and "double-backs" using bushes, wind-fallen trees, stumps 
and many others. Their ways of hunting, as shown above, are highly 
variable and they are usually governed by the complex and highly 
coordinated behavior of the members of the litter (pack). The complex 
nervous activity of wolves with the highly developed and multiple 
conditional reflexes (temporal connections) allows them to adapt them- 



245 

selves quickly to new situations. This "flexibility" in the behavior of 
wolves, for example, their ability rapidly to distinguish poisoned baits 
and avoid them, makes their control difficult. 

Seasonal migration and transgressions. Wolves in the majority of 
regions are settled. They abandon their accustomed places quite unwill- 
177 ingly, usually only in cases of severe shortage of food in their hunting 
territory. The migration of wolves is, in the majority of cases, con- 
nected with nutrition and represents nomadic movements for food. 

Significant seasonal movements of wolves have a place only in 
tundra, steppes, deserts and montane regions with nomadic (shifting) 
livestock breeding, where the herds of reindeer, horses and cattle are 
regularly driven from the summer pastures to winter and vice versa. 
Examples of such nomadism are performed by herds of reindeer in 
tundra, saigas, dzerens and goitered gazelles in steppes and deserts, roe 
deer in the Urals and in Siberia, and by deer in the mountains. The 
times and paths of migration of the ungulates and the wolves coincide. 
In mountains, during the fall of deep snow, wolves, following the 
ungulates, move to less snowy places, usually descending to the foot- 
hills. 

In the tundra wolves lead a nomadic way of life for three-fourth of 
the year, not adhering to stable hunting regions. Twice yearly they 
accomplish large meridional migrations connected with the driving of 
the [domestic] deer herds. When the deer herders shift southward, the 
wolves go together with the deer herds to the forest border. They do not, 
as a rule, penetrate deeply into the taiga, although in the European north 
in particular, the main mass of the deer herds of the state farms and 
collective farms winter there. Wolves prefer to remain in the forest- 
tundra and mossy bogs with less deep and more firm snow cover. Here 
in the forest-tundra, there is much natural food, since willow ptarmigan 
migrate thither, white hares are very frequent, and until recently, elk 
were encountered along the river valleys where trees are located 
(Semenov, 1954). In the Nenetsk Nats. Region, besides seasonal migra- 
tions, the movements of wolf packs from the Bol'shezemel'sk to 
Malozemel'sk tundra in connection with herd migrations were noted, 
but return movements were not (Makridin, 1959). Seasonal migrations 
of tundra wolves in the European north reach 200-300 km (Semenov, 
1954). 

A small portion of the wolves remains on the tundra in winter. 
They move out to the sea coast, where remain only small reindeer herds 
of hunters engaged in the arctic fox harvest and near fishery stations, 



246 

feeding on fish and other animal wastes. On the seashore wolves find 
the bodies of stranded marine animals. Tundra wolves are very mobile 
and move for great distances. A trailed pair of wolves went 70 km in 
one night (Semenov, 1954); from an а1ф1апе it was possible to follow 
the path of a pack of wolves that went more than 150 km in 20 hours 
(Semenov, 1954). Regular migrations to the Novosibirsk islands, fol- 
lowing reindeer across the sea ice are accomplished by wolves in 
northern Yakutiya. In the forest zone, wolves are less mobile. They 
follow roe deer during their migration in the taiga of eastern Siberia 
(Khudyakov, 1937). During food abundance, the predators remain for 
5-8 days in comparatively small territories; if they do not obtain food, 
the wolves go "on foot", moving 25—50 km in a 24-hr period. 

In the montane forest landscapes of the southern Urals, wolves 
migrate with the ungulates in winter from places of deep snow. In 
Bashkir preserve wolves, together with roe deer and marals, remain 
in the winter in the montane massif of South Krak and stay in small 
numbers in the Uraltau (Kirikov, 1952). In the Altai, they leave the 
mountains following the roe deer into the lower valleys. In the Tien 
Shan, the predators ascend in summer into the mountains (to high 
watersheds) following the herds of wild ungulates and domestic ani- 
mals and they again descend lower when deep snow falls in the 
mountains. 

Generally, settled populations of wolves live in Caucasian, 
Borzhomsk and Zakatalsk preserves, as well as in the reed beds of the 
Caspian coast; however, they also perform vertical and horizontal mi- 
grations following the wild ungulates. In the remaining territory of the 
Caucasus, especially in its eastern part, wolves wander after the herds 
of domestic livestock, ascending the mountains in summer and in 
winter — ^to the lowlands (Vereshchagin, 1959). 

Wolves accomplish regular migrations in the steppe regions, where 
transhumant animal breeding exists or where large herds of wild ungu- 
178 lates survive. Following herds of dzeren [Mongolian gazelle] on the 
[USSR] boundary in the Trans-Baikal steppe, wolves frequently pen- 
etrate from the Mongolian Republic. In the deserts of central Kazakhstan, 
these predators accompany the herds of saigas and goitered gazelles 
going, in September-October, to wintering grounds in the southern part 
of Betpak-Dala. From the Severo-Kazakhstansk district, the majority of 
wolves, in December— April, leave for more southerly regions (Afanas'ev 
et al., 1953). In the Kustanaisk steppes, in the periods when wolves 
were abundant in 1920—1921, in autumn they moved about, following 



247 

the herds of horses and going by day behind the horse drov- 
ers at a distance of only few hundreds of meters or even nearer (V.G. 
Heptnef). 

In deserts and in part in the steppes, the cause of wolf migrations 
may be steppe fires, drought and lack of water or other reasons leading 
to massive translocation of livestock and the migration of wild ungu- 
lates. In connection with fires in the Badkhyz desert and consequent 
lack of water, goitered gazelles appeared en masse along the Kushka 
river in June 1942. Following them thither, the wolves appeared. This 
massive appearance of gazelles, and of the wolves following them was 
noted in 1927 along the Murgab river. In other desert-steppe regions 
of such a sort, migration of desert and steppe ungulates to waterholes, 
in the middle of summer chiefly in the river valleys, are always accom- 
panied by corresponding migrations of wolves (Heptner, 1956). 

Besides migrations for food, usually taking the character of sea- 
sonal migrations, wolves sometimes perform long, time-consuming 
massive migrations not having a regular character. They are either 
connected with unfavorable changes in their principal habitats, or rep- 
resent the growing cubs establishing themselves. 

In some years, it was noticed that tundra wolves migrate in a 
horizontal direction. Migration of wolves was observed from the Urals, 
to the Yamal, into the Bol'shezemel'sk tundra; the wolf packs crossed 
the Malozemel'sk and Timansk tundras and reached the Kanin isthmus. 
Such migrations of wolves usually coincide with massive migrations of 
arctic foxes in that direction. A return movement of wolves (from west 
to east) was not observed (Semenov, 1954). 

In Kazakhstan in individual years wolves also perform long and 
irregular movements. Thus, in January 1941 in Dzhambulsk district 
many "Siberian" wolves appeared, which were well distinguished from 
local ones by their large size and gray color. In autumn 1942, wolves 
moved en masse from the northern Caucasus, from Rostovsk and 
Volgogradsk districts into Zapadno-Kazakhstansk, Gur'evsk and even 
Aktyubinsk districts of Kazakhstan. The large European wolves well 
differentiated from the small, reddish local ones (Afanas'ev et al., 1953). 

Settlement of wolves is easily registered when they enter into places 
which are unsatisfactory as habitat. Thus, during the summer, wolves 
penetrate into the taiga of western Siberia from districts comparatively 
far to the north and south, that are continuously inhabited, for tens or 
even hundreds of kilometers (Aleksandrovsk region of Tomsk district, 
upper reaches of the Taz river, to Surgutsk region; Leptev, 1958). 



248 

However, when the deep snow falls, they abandon these places or die 
there. Wolves which in winter inhabit the Berezovskii region of 
Tymnensk district along the Syn' and Lyapin rivers, migrate in summer 
to the Urals (Flerov, 1933). 

Under difficult conditions lengthy migrations are known. Thus, 
from 1933 to 1945, wolves four times penetrated over ice to Barsa- 
Kel'mes Island in the Aral Sea which hes 80 km from the shore 
(Afanas'ev et al., 1953). In 1938, three wolves walked on ice about 
200 km and reached Wrangel Island. 

Reproduction. Wolves are monogamous; the united couple usually 
stays together for life, if one of the wolves does not die. Upon the death 
of the male or female, as shown above, pairs are quite quickly re- 
established. Since in wolf populations, males often predominate (Teplov, 
1948), unpaired females are a rare occurrence, while unpaired males 
are quite common; this is related to the later maturity of males. Among 
179 wolves caught in traps and from aiфlanes in the European tundras, 
71% were males in 1951-1953 (31 wolves studied); in 1954, 67% 
males (30 wolves) and in 1955, 50% (72 wolves); the average of all 
years (133 wolves) was 59% (Makridin, 1959). 

Wolves mature at about 2 years; practically all she-wolves become 
sexually mature at the end of the second to beginning of the third year 
of life; males in part begin to participate in reproduction in the third, 
and in part the fourth year of life. Estrus in she-wolves and rut begin 
usually at the beginning to middle of the second half of the winter. Old, 
multiparous she-wolves enter estrus first, approximately 2—3 weeks 
earlier than heat in young, just maturing she-wolves. Before the begin- 
ning of rut, as was shown, the wolf pack dissolves. Adults disperse first, 
then yearlings, and finally juvenile wolves. Later on, they re-unite, but 
they do not approach the adults and lie separately. Driving away the 
young, the adult male becomes unusually attentive to the she-wolf, does 
not leave her for a minute and opens up a path for her in the snow. 

Heat in older animals usually proceeds calmly, but in certain (rare) 
cases it becomes complicated due to the presence of 1—2 unpaired 
males. Heat lasts approximately 2 weeks. At the beginning of heat and 
rut in the young wolves, they drive away the juveniles still living with 
them at that time. Particularly at this time, during the formation of new 
pairs, the so-called "wolf wedding" is manifested, when a group of 
males gathers around one female. These "weddings" are accompanied 
by violent struggles and hard fights by young males for the female. The 
fight sometimes ends with death and the dead wolf may even be 



249 

devoured. Such a group does not remain in one place for long, but 
wanders widely spending the day in remote places — ^wilderness, bogs, 
fir-groves, often wandering the byways, usually at night but sometimes 
also by day time. The she-wolf bonds with the strongest wolf of this 
pack and thus a new pair is formed, lasting until the death of one or 
both "spouses." 

The periods of estrus and rut vary in different regions. In the south, 
copulation takes place 1-1 .5 months earlier than in the north; therefore, 
the period extends from December to' March. In the Karelian ASSR, 
estrus is initiated at the end of February and in March (Marvin, 1959). 
In Arkhangel'sk dist[rict], rut begins at the beginning of March in the 
northern regions, and at the end of February in the southern, while in 
regions adjacent to Vologodsk district, it begins approximately one 
week earlier (Semenov, 1954). In the forest-tundra and tundra of Nenetsk 
Nats. Region, the first female in which ovulation had taken place was 
caught on the 9th of March. In the period 9 March— 14 April, only 
yearling females were still not ready for the rut (Makridin, 1959). In 
Tatariya, rut usually occurs in the middle of February. In the middle 
Urals, wolves copulate in February (Pavlinin, Shvarts, Danilov, 1951). 
In forested montane landscapes of the southern Urals, estrus in wolves 
proceeds from the end of February-beginning of March (Kirikov, 1952). 
In the former Tarsk region, estrus in wolves proceeds from the second 
half of January— first half of February, and whelping in April (Shukhov, 
1928; Kuklin, 1938). In Irkutsk district, estrus begins in wolves at the 
end of January (Timefeev, 1949). 

In Belorussia, rut is initiated at the end of January; it ends approxi- 
mately at the middle of March (Serzhanin, 1955). In Voronezh district, 
rut in adults begins at the first, or more often the second third of 
January. At this time, the characteristic howl of wolves can be heard 
around dawn, but two weeks later, the breeding pack of one female and 
2-3 (here rarely more) male wolves can be found during day. Rut ends 
at the beginning of February, but sometimes extends to the first half of 
March. After that, the formed pairs occupy territories, and, in April, the 
she-wolves give birth (Mertts, 1953; Barabash-Nikiforov, 1957). In 
Ahna-Atinsk district estrus in she-wolves proceeds from December- 
January. In the lower Hi river, a she-wolf in a state of estrus was 
caught on 27 January 1949. In northern Kazakhstan, estrus occurs in 
February-first half of March. 
180 The duration of pregnancy in the she-wolf is 62-75 days. 

In Nenetsk Nats. Region, the largest number of embryos occurring 



250 



in pregnant she-wolves was 9; the average number of embryos was 6.5 
per female. From one to nine pups have been found (6.7 per female on 
the average) (Makridin, 1959). The number of whelps in a wolf litter 
in Yamalo-Nenetsk Nats. Region is more often 3-4, rarely 5. The usual 
number of a pack (litter) is 7-9 wolves, including 2-3 yearlings 







!йГ ^Ж- As 



"«ft 






Fig. 68. Utems of she-wolf with 9 embryos and uterus of a young nulliparous 
she-wolf, [both] caught in March. Photograph by V.P. Makridin. 



251 

(Rakhmanin, 1959). In the middle Urals, there are up to 12 young (up 
to 14 fetuses were found), more frequently 5—8 (Pavlinin, Shvarts, 
Danilov, 1951). Young she-wolves produce smaller litters, usually 3— 
5 young; adults are more fecund (6-8 and up to 14 young). Old females 
in Belorussia produce 8 young, rarely more, and young — 3—5, giving 
an average litter size of 4— 6 pups (see Tables 16 and 17). 

The average size of litters varies in different years. 

Before whelping, the she-wolf usually returns to the old den in 
which it has raised the past litter. A young one, whelping for the first 
time, searches for a new den, usually not far from its birth place. 

Parturition of pups takes place in the warm months, when the 
stocks increase and food becomes varied. In Karelia, 3-7 (rarely more) 
blind pups are bom in April-May or the beginning of June (Marvin, 
1959). In Arkhangel'sk district, forest wolves whelp at the end of April 
181 and the first third of May. They give birth to 3-12 pups (more often 
5-8). In the tundra, pups are bom around the second half of May- 
beginning of June; the young are from 3 to 8, 6 on the average (Semenov, 
1954). In the southem Urals, pups appear at the beginning of May. On 
21 April 1946 a killed she-wolf had embryos of about 310 gm each 
(Kirikov, 1952). The earliest litters are found in Belomssia in the 
second half of March, more often in April and May. In Brestsk district 
in 1948, pups still blind were found on the 10th, 22nd and 27th of May. 

In the central chemozem districts, there are usually 3^ pups in the 
litter. In 1950 litters of 6 and 8 pups were captured, and in 1956 a litter 
of 10 pups. The low average [3^] litter size in these districts can be 
explained by the fact that the majority of females die when they are still 
young, far from reaching the age for which a high litter size is char- 
acteristic (Mertts, 1953). 

In the Ukraine the pups are bom at the middle of March-beginning 
of April. The earliest litter was found on 3 January 1949, of 5 one- 
week-old pups (Kiev district). In Sumsk district, two pups approximately 
3 weeks old were found on 2 June 1947. In the first third of June, a 

180 Table 16. Frequency of occurrence of wolf litters of various sizes in 

Belorussia (Serzlianin, 1955) 



Number of i 2 3 4 5 6 7 8 910III2 Total 

pups in litter 



Number of 5 3 4 10 14 13 6 7 — I — 1 65^ 

litters 



♦Should total 64 — Sci. Ed. 



252 

Table 17. Average number of pups in a den (Serzhanin, 1955) 

Years 1947 1948 1949 1950* 1951 1952 



Number of dens 


81 


186 


150 


168 


180 


173 


Average number of 


4.5 


4 


4.5 


5.2 


4.9 


4.2 


wolf cubs in den 















*In Russian original, 1959 — Sci. Ed. 

Table 18. Frequency of occurrence of wolf litters with different numbers of 
young in Kazakhstan (Afanas'ev et al., 1953) 



Number of pups 
in the litter 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


12 


13 Total 


Total number 
of cases 


2 


3 


7 


10 


9 


12 


5 


8 


2 


5 


2 


1 66 



Including: 
In southern 

districts — — 2 2 3 3— 2— — — — 12 

In northern and 

central districts 235869562521 54 



litter was found in Chernigov district, and the latest litter was found in 
Volynsk district in July 1949 (Komeev, 1950). In Zakaфatsk; district 
with its mountainous climate, in the majority of cases birth of the young 
usually takes place at the end of May. In individual cases, in the 
Ukraine, litters attain 10-11 young. 

In the north and the central districts of Kazakhstan, small pups 
were found on 16 April (2 litters), from 2 to 30 May (17 litters) and 
in June (1 litter). From 2 to 13 pups occur in litters, the average litter 
size in the southern districts of Kazakhstan being 6.3 pups and in 
northern and central districts, 7.0 cubs (Table 18). A Utter of 13 pups 
was found once, the mother and pups being in extreme emaciation 
(Afanas'ev et al., 1953). 

In southwestern Turkmeniya, the she-wolf produces young at the 
end of March-beginning of April; on 12 April 1942, blind pups 5-7 
182 days old were caught. In the litter 3-4 pups usually occur (Dement'ev, 
1959). In the Caucasus, the she-wolf gives birth to 5-6 pups on the 
average. Time of birth of pups extends from March to May (Satunin, 
1945). In Azerbaidzhan, litters appear in April— May (from 2 to 8 
young) in secluded places — remote ravines and in dry islets in marshes 
(Vereshchagin, 1951). In Armenia, wolves copulate in January, giving 
birth to cubs in March; the litter is 3-8 young (Dal', 1954). 



253 



(^f^(BK5l.ra5i!S?KHPi^S^S^^'.- ■ 



Fig. 69. Wolf pups 10-11 days old, taken away from the den. Khakassiya, 
Krasnoyarsk territory. 18 May 1956. Photograph by V.V. Kozlov. 

Maternal sense of wolves is very strongly expressed. A case is 
known when a she-wolf followed a hunter who had captured its cubs 
into a village and at night walked near his house for a long time. 

Out of 5—7 pups (average litter size) bom, only 1—3 animals reach 
sexual maturity (juvenile mortality is 60-80%). Wolves usually live 
not more than 15 years. A wolf which lived for 16 years in captivity 
was quite decrepit (Afanas'ev et al., 1953). 

Growth, development, and molt. Wolf pups are bom blind, with 
closed ear openings, and covered with short soft grayish-brown fur. 
The weight of new-boms is 300-500 gm. They begin to see on the 9th— 
12th day; at the end of the second — beginning of the 4th week, milk 
canines erupt. In captivity, pups which begin to see on the 9th— 14th day 
grow strong and healthy, while those in which vision develops later 
remain usually sickly and ricketic. 3-week-old pups crawl out of the 
den, and at 1.5 months old, they quickly flee from danger and take 
cover. Adult wolves leave their 3-week-old cubs alone when setting 
out on a hunt. 

In the first weeks, the mother does not leave the litter and during this 
time the father-wolf feeds her, bringing prey to the den, or regurgitating 



254 

the semi-digested meat. The she-wolf nourishes the young with milk 
until they are 1.5 months old. When the pups are 3^ weeks old, they 
begin to eat the regurgitate offered by the father, and the mother only 
eats the remains. The necessity of regurgitative feeding (semi-di- 
183 gested meat) is, apparently, related to deficiencies in digestive juices 
(peptidases) in the young whelps. Artificially fed pups without such 
regurgitate are accompanied by poor growth, delayed development 
and rickets. Pups 3-4 months of age already find such regurgitate 
unnecessary. In summer, nursing she-wolves become very thin. Milk 
gradually becomes, for the young, only a supplementary food, and 
they feed mainly on small animals which the father and mother bring 
them. 

In the first four months, the tempo of growth of the pups is high. 
During this time, their weight increases nearly 30 times (from 350- 
450 gm to 14-15 kg; Novikov, 1956). In the southern Urals, the 
weight of a juvenile male in August 1941 was 1 1 kg, of a juvenile 
female — 8 kg, in September — 13 kg. In the next 3^ months, the 
weight of wolves doubles and in December, they reach 28—30 kg 
(Kirikov, 1952; Gavrin and Donaurov, 1954). In Ryazansk district, 
young at 6 months attain a weight of 16-17 kg. 

Later, when the pups become strong, the mother and father bring 
living, but injured, prey to them in the den, teaching the whelps how 
to catch and kill it. At the middle of summer, the adults begin to lead 
the pups to killed prey. The actual training of the young begins at the 
second half of the summer. From August, they attempt to catch rodents 
and other small animals by themselves, and starting from September 
they share with the adults in hunting ungulates. 

The molt of wolves takes place twice per year; in spring from the 
end of March or middle of April to July. In autumn there is no full 
replacement of the pelage, only the underfur growing and developing 
(from the beginning of September to the end of October, and sometimes 
to the end of November). 

Enemies, diseases, parasites, mortality, competitors. This is little 
studied. The ticks Ixodes ricinus, Dermacentor pictus, Sarcoptes scabiei 
[the mange mite] and others parasitize wolves in great numbers. In 
distinction from foxes, only a little full-blown mange is ever observed 
on wolves; it more often strikes young wolves. Among other ectopara- 
sites may be noted: the biting louses Trichodectes canis, the sucking 
louse Linognathus (sp.), and the fleas Pulex irritans and 
Ctenocephalides canis (Komeev, 1950). It is completely obvious that 
this list is very incomplete. 



255 

Among endoparasites are established the nematodes Toxocara 
leonina, T. canis, the cestodes Taenia pisiformis , T. hydatigena, Echi- 
nococcus granulosus, Mesocestoidea lineatus, Dioctophyme renale 
(through fish) and the adult phase of the causative agent of sheep 
gid[diness] — Multiceps multiceps. Infections with helminths are not 
great. In the central chernozem districts, of 20 wolves examined, nema- 
todes were found in the stomachs of seven (4-1 1 specimens) in the 
small intestine of one — many fine cestodes, and in the trachea — 5 
nematodes (Barabash-Nikiforov and Pavlovskii, 1947). 

Wolves frequently suffer from rabies and they act as a source of 
this infection for people. The overwhelming majority of "attacks" of 
wolves on humans stem from rabid wolves which, because of the dis- 
ease, lose the caution and fear of man of normal wolves. Although 
70—88% of the total cases of rabies among humans is due to bites by 
dogs, cats — 6-8%, and wolves — ^less than 2—3% (Rudnev, 1950), wolves 
play the role of principal host (reservoir) of the rabies virus in nature 
(Zhdanov et al., 1955). 

The only serious enemy and competitor of wolf is, apparently, the 
tiger. In the Ussuri territory, wolves were absent in some places from 
time immemorial and they appeared only after tigers were extermi- 
nated. It was also noticed that wolves disappear from those places in 
which tigers appear. 

The mortality of wolves is great. In the course of the first year of 
life, as shown, it reaches 60—80%, and subsequently decreases to 
approximately 40—50% per year. Because of this, young of the cur- 
rent year usually compose about one third of the population (Makridin, 
1959). 

Population dynamics. The number of wolves, though relatively 
stable, changes under the influence of different conditions of existence. 
184 It is necessary to accord the first place for significance their pursuit by 
humans, which increases with increase in density of settlement and the 
development of agriculture; it becomes weakened when agricultural 
activity is disturbed for one reason or another. Therefore, a rise in the 
number of wolves was observed after each war, famine or economic 
dislocation. 

At the beginning of the nineteenth century, wolves were numerous 
in the majority of the developed regions of the country, including the 
entire Ukraine both in Poles 'e, and in the steppes of the Black and 
Azov seas. The abundance of predators in the forested parts of the 
Ukraine was related to the abundance of wild mammals (deer, roe deer. 



256 

wild boar, beaver, hares) and birds. The number of wolves particularly 
increased after the French invasion in 1812 when the numerous bodies 
along the route of the French retreat served as food for wolves. At that 
time, there was even noted a migration of predators to Ukrainian Poles'e 
from neighboring districts of Belorussia (Komeev, 1950). At the middle 
of the nineteenth century, although the populated areas in the Ukraine 
increased significantly, wolves remained numerous even in the vicinity 
of such large cities as Odessa. In only 4 days in the autumn of 1 844 
during a mass drive in [the former] Khersonsk, Ekaterinoslavsk, 
Tavrichesk and Bessarabsk governances, 1404 wolves were caught 
(Brauner, 1923). In the eighties of the past century, there were many 
wolves in Chernigov governance (Dubyanskii, 1885). 

At the end of the nineteenth century, the number of wolves in the 
entire middle zone and in the Ukraine noticeably decreased. At the 
beginning of the twentieth century the wolves remained in significant 
numbers only in districts with large tracts of forests and sparse popu- 
lations. In [the former] Khersonsk, Nikolaevsk and Odessa provinces, 
they were completely destroyed, and in the other southern districts 
wolves remained only in the floodplain of the Danube (Izmail'sk dis- 
trict). At the time of the First World War, the number of wolves grew 
shaфly; this was enhanced by the Civil War and the devastation ac- 
companying it. By 1920, the wolves occupied nearly all of the Ukraine, 
central chernozem district, the middle non-chernozem zone and the 
Povolzh'e. In Chernigov district in 1923 alone, wolves destroyed 5.2% 
of the domestic livestock count. As was noted, wolves were, at that 
time, very numerous in the north of Kustanaisk district (V.G. Heptner). 

In subsequent times, the number of wolves began to decrease and 
at the beginning of the Great Patriotic War [W.W. II] wolves de- 
creased overall by approximately 100 times as compared to 1920. At 
this time in the Ukraine, wolves remained only in Zhitomirsk, the 
southern regions of Kievsk and Chemigovsk, and southwestern regions 
of Khar'kovsk districts, in the Starobel'sk steppe and in Lugansk dis- 
trict (Migulin, 1938; Komeev, 1950). 

New proliferation and recolonization of wolves took place in the 
years of the Patriotic War, 1941—1945. The number of wolves grew 
intensively in places where in addition to food abundance, wolves had 
well protected places for constructing dens. After the Great Patriotic War, 
the number of wolves increased in all districts without exception, and 
they appeared in the places where they had been previously exterminated. 
In the Caucasus, where the number of wolves had increased somewhat 



257 

after 1927—1930 and remained at quite a high level until the Second World 
War, it shaфly increased in the period 1941-1945. Only after 1950, as a 
result of the strengthening of wolf control did their numbers in the Caucasus 
noticeably decrease (Vereshchagin, 1959). 

Thus also proceeded [events] in the European north and in western 
and eastern Siberia. The distribution of wolves in Siberia was favored 
by winters with little snow at the end of the 1940's. Wolves became 
the scourge of [rein] deer breeding in the Katangsk region of eastern 
Siberia and appeared in the Shitkinsk and Bodaibinsk regions (Timofeev, 
1949). 

After the war, wolf control was shaфly intensified, and their num- 
ber quickly began to decrease everywhere. This was favored by the 
185 application of new, totally destructive methods of hunting (see below). 
In Arkhangel'sk district, intensive destruction of wolves stopped their 
growth in numbers, and then shaфiy reduced them. The number of 
captures per year in Arkhangel'sk district increased from 129 in 1948 
to 360 in 1953 (in all 1374 wolves were destroyed in 1948-1953; 
Semenov, 1954). 

In addition to this basic factor in the population dynamics of wolves, 
great significance must be placed on nutritional conditions in winter 
and the supply of water in summer. The first is of basic importance in 
the forest zone and the second, in the steppe and desert zones. How- 
ever, in the steppe regions of Kazakhstan, the number of wolves also 
shaфly decreases in winters with much snow, mainly as a result of 
death from starvation. In the winter of 1948/49, which was character- 
ized by strong and prolonged blizzards, wolves were not able to capture 
food. They gathered near the settled points and somehow fed on dogs, 
but they quickly began to die (Afanas'ev et al., 1953). At the same 
time, the widespread opinion that all wolves are obliged to fast in 
winter is not true. In a pack (family) of two grown-up wolves and four 
juveniles caught in December 1940 in Moscovsk district (near 
Maloyaroslavts), the adults were well nourished, but not fat, while the 
young had good deposits of internal fat which nearly covered the kidneys 
and a little fat on the thighs. It is evident that the old wolves, even in winter, 
were solicitous about the better nutrition of the young. Accessible livestock 
are not here in winter, and only carrion was at the disposal of the pack, 
apparently, elk, hares and dogs being scarce in the district (in the stomach 
of one wolf, a piece of leather collar was found; Heptner and Morozova- 
Turova, 1951), Communications about the capture, in winter, of fat and 
"very fat" wolves are not very rare, particularly from the tundra 
(Makridin, 1959). 



258 



., И 







^»к 



./ V 

Pig. 70. Tracks of a pacing yearling female. Oka preserve. Ryazansk district. 
15 January 1949. Photograph by V.G. Heptner. 

After a series of drought years in semi-desert and desert regions, 
the number of wolves shaфly decreases. Thus, in 1939 in the Ustyurt 
and Mangyshlak, wolves almost totally disappeared, and in the Turgaisk 
region of Kustaniask district in 1938, the number of predators shaфly 
1 86 decreased. The drying-out there of steppe springs and lakes deprived 
the wolves of waterholes and the surviving animals concentrated m 
abundance in ravines at the village of Borgot. In the bottom of ravines, 
the wolves dug holes, in which ground water gathered, and the animals 
came regularly to water at these "excavations" (Afanas'ev et al., 1953). 



259 

In the drought of 1936, all wolf litters living in the lower Sarysu river 
and adjacent regions, gathered in small reeds surrounding the two re- 
maining TelekuF lakes. They attacked everything available for them to 
pursue, but they apparently were starving and the young grew poorly 
(Sludski, 1937). 

Among wolves epizootics sometimes develop, but their causes, 
course and after-effects are practically unstudied. 



iMg. 71. Tracks of pacing wolf. Krasnoyarsk territory. December 1959 
Photograph by V.V. Kozlov. 



260 

The reproduction of wolves in different years is not the same. In the 
central chernozem districts, intensive reproduction was observed in 1925, 
1934 and 1935, in 1943, 1944 and 1947 (Barabash-Nikiforov, 1947). 

Field characteristics. The wolf is a strong, enduring animal; with- 
out special effort it can pull or carry on its back a ram. It can move 
quickly, achieving a speed on bare ground* of 50-60 km per hour 
(Semenov, 1954). Information about speeds up to 80 km per hour 
(Nanteifel', 1948) is completely unverified. Usually wolves run at a 
trot. This trot nonetheless allows them to cover great distances. In this 
gait, the track of the hind foot falls exactly on the track of the fore, and 
the tracks are situated, as along a string in one line. During walking, 
this exactness in the position of the feet is not observed. Wolves of a 
well-fed pack go in single file, and a tired lead animal is replaced 
periodically. This allows the pack to move for long distances even in 
deep snow. Hungry wolves sometimes separate from each other. 

When running, the wolf holds its head somewhat low and a little 
to the side, directing one ear forward and the other backward. This pose 
gives them the possibility of using their exceptionally shaф hearing. 
Changing the gait when necessary, the animals switch to a gallop and 
then a walk; they often stop and stamp in place. In deep snow, they 
187 prefer to use roads. By day they go at a walk, avoiding populated 
places; sometimes dodging and doubling back, like a hare, usually 
shortly before lying down. Alarmed, wolves may go for 70 km in one 
night at a trot or jog (Komeev, 1950). Wolves can swim well. In 
autumn 1948, a litter of wolves regularly swam across the Desna river, 
on setting out to hunt and on returning from it to the den until a strong 
cold snap [occurred] (Komeev, 1950). In Ryazansk district in the 1950's, 
in periods of night hunting, they swam across the swift, wide Oka river 
to attack the livestock on an island (V.G. Heptner). It is very difficult 
to catch up with untired animals by rowboat. 

The tracks of wolves resemble those of dogs, but differ in their 
form and large size. The wolf does not "spread" the digits in the track, 
but holds the pads, as hunters say, more "balled up". The print of the 
anterior pads of a large central Russian wolf reaches 123 mm in length 
and 55 mm in breadth. The track of one wolf in the Priozem region of 
Arkhangel'sk district had a length of 130 mm and a breadth of 97 mm. 
The track of a huge animal, killed in Vladimirsk district in 1953, was 
15 X 12 cm in shallow freshly fallen snow (N.D. Sysoev). The largest 

*Literally chernotropa, or "black path;" see p. 242 — Sci. Ed. 



261 

track of the anterior pad of a tundra wolf is 126 x 92 cm, and of the 
posterior, 123 X 80 mm (Timansk tundra; Semenov, 1954). The soles 
of the pads of the tundra wolf are better furred and the toes are more 
movable (expanding more widely). In tracks, females and males can be 
differentiated according to the manner of urination (the first is directly 
on the tracks, the second to the side). The she-wolf and the young leave 
their feces directly on the tracks, whereas the wolf usually moves aside 
from the tracks. 

Trails of wolves differ from individual tracks in the somewhat 
larger size of the tracks, their compactness, the wider "drag" — the 
furrow in the snow, inscribed by the end of the wolf pad in the begin- 
ning of the step, and the "pillow case" — such a furrow at the end of 
the step (in front of the foot print). 



Fig. 72. Wolf trail. Track of four wolves proceeding "track in track". Khakassiya, 
Krasnoyarsk territory. December 1 956. Photograph by V.V. Kozlov. 



262 

The voice of a mature wolf is a long bass howling. The male wolf 
gives voice through an octave, passing to a deep bass with stress on 
"O", and the she-wolf produces a modulated baritone through the nose, 
with stress on "U". Her voice is particularly mournful and wild. Yearlings 
howl with characteristic yelps resembling those of dogs. They include 
high notes and end by a characteristic barking. Juveniles almost never 
howl, but whine with laugh-like, thin voices. Howling wolves are heard 
for a distance of up to 2 km. The meaning of the howl is an assembly 
call for the mature wolves returning with prey and a signal of well- 
being in the region of the den. The male howls first, the female answers, 

188 and the young repeat in a dissonant chorus. Howling begins at twilight 
after sunset, and continues with pauses of usually not more than 1 5 
minutes. Wolves do not answer a howl in rainy weather and when 
satiated (Danilov, 1945). In autumn and spring (until the birth of young), 
and sometimes in winter, wolves usually howl when going to hunt. In 
certain periods, for example, in the forests of the lower Samur river in 
southern Dagestan in August 1924, they "struck up a tune" with great 
precision at the coming of darkness (V.G. Heptner) (N.N.). 

189 Practical Signiflcance 

Wolves are harmful to humans in many respects: they attack livestock 
and dogs, wild ungulates and other useful animals, spread diseases and 
attack people directly. The little use which may be derived from cap- 
tured wolves (skin, tasty meat which is fully suitable for food) as well 
as sporting pleasure of the hunting of wolves, are not to be compared 
with the damage to human health and economic interests caused by 
these undoubtedly injurious predators. 

The average agricultural losses caused by wolves in the middle of 
the past century in only the 45 European governances of Russia were 
estimated at 179 thousand head of large homed livestock and 563 
thousand of small annually, for a value of 7 million 600 thousand 
rubles in those day's prices (Lazarevskii, 1976). Animal husbandry in 
Vologodsk, Novgorodsk, Minsk, Volynsk, Kaluzhsk, Orlovsk, Ryazansk 
and Samarsk governances suffered especially from wolves. In 1871— 
1873, wolves killed an average of 2550 (0.29%) head of large homed 
livestock and 8550 (0.65%) of small livestock per year in Kiev gov- 
emance alone. In Volynsk govemance at that time, mortality reached 
1.36% per year (12,300 head of large homed livestock and 19,600 of 
small; Komeev, 1950). Recently, the amount of harm sometimes 



263 



Wolf ^Cc »Ct <t Kr Kt »& Ks »r= 



»? 



Dog 



•- .u •^ 



»5t 



•& 



Wolf K? 



«1= 



•& 



Ks 



Dog »S 



»& 



♦Is 

2 



K' 



WoH 



»fs 



»& 



•& 



•& 



♦:- 



»& 



♦jt 



K= 



Wolf 



»:=: 



<- 



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Fig. 73. Scheme of tracks of wolf and dog in different paces of running (from 
Kozlov, 1955). 1 — walking slowly, 2 — trotting, 3 — pacing, 4 — galloping. 



'^Шш)- 



188 Fig. 74. Prints of front and hind feet of wolf on soft silt. Ganyshkino, in the delta 
of the Volga. 23 October 1951. Sketch by A.N. Formozov, about 1/2 natural size. 



264 

decreased and sometimes increased, growing particularly during social 
and economic shocks (wars, economic crises, crop failure and so on) 
as a result of weakening of predator control. This was noticed both with 
us, and abroad as well. 

The harm caused by wolves greatly increased due to growth of 
their number after the First World War and then in the Civil War. In 
three republics: the Russian Federation, the Ukraine, and Uzbekistan — 
865 thousand head of livestock were registered as killed by predators 
in the agricultural year 1924/25, a sum of 15,530 thousand rubles in 
the price at that time. Approximately 35% of the losses took place in 
Siberia and Kazakhstan, followed by the other main stock-raising dis- 
tricts. In relation to all livestock in general, these losses consisted, in 
the lower Volga of 2.2%, in Siberia — 1.6%, in Kazakhstan — 1.5%, 
and for the whole territory of the USSR, 0.54% on average. Among the 
animals killed, 16% were horses (4.2% adults, 1 1.8% young), 18.1%— 
large homed livestock (6.8% adults, 11.3% young), 52.6% — sheep, 
5.8%— goats and 7.2% — swine (Krasil'nikov, 1926). About 30%) of 
the animals were killed in winter in the time of confinement in struc- 
tures or in winter quarters. Many domestic poultry and dogs are killed 
by wolves. Hundreds of thousands of geese were exterminated yearly. 
In summer, they sometimes even comprised the principal food of wolves 
in a series of places. Although losses were reduced in 1928 due to the 
return of wolf control, until 1941 they remained in the vicinity of 
several hundred thousand head of livestock annually, i.e., quite high. 
In connection with growth in the number of wolves at the time of 
the Great Patriotic War and afterward, there was a shaф increase in 
damage caused by them. Thus, in 1944 in the Buryat ASSR, more than 
5300 head of livestock were destroyed by wolves; in Penzensk district, 
780 horses and cattle, more than 5000 sheep, 300 swine and others 
were killed in 10 months of that year. Losses for the whole year 
approximated 8700 head of livestock in all districts. In Kuibyshevsk 
district, this figure reached 4200 head of livestock. In sheep-raising 
state farms, the wolves killed at that time 1-1.5%) of the breeding 
flocks each year (Danilov, 1945). In Vomezh district, only the wolf 
population connected with Usmansk forest caused losses in the forest 
itself, and on 295 collective farms situated in its neighborhood, esti- 
mated at 760 thousand rubles in four years (1946-1949). During that 
time wolves killed on collective farms 164 calves, 163 foals, 3119 
goats and sheep, 37 pigs, 3174 dogs, and 2500 geese. At that time as 
190 well, in Usmansk forest wolves destroyed 200 deer and not less than 



265 

19 beavers (Martts, 1953; Barabash-Nikiforov, 1957). In 1946 in the 
Zyryanovsk and Asinovsk regions of Tomsk district, 600 head of live- 
stock were slaughtered by wolves (Boikov, 1949). 

The amount of damage caused by one litter of wolves can be 
judged by the fact that in Ryazansk district (Bel'kovsk region) 2 ma- 
ture and 6 young wolves destroyed during 8 months (from May to 
December 1945) 6 horses, 1 foal, 5 cows, 150 sheep and goats, 3 pigs 
and many domestic geese, ducks and hens (Kozlov, 1955). 

Annual losses to reindeer-breeding caused by wolves in the Yamalo- 
Nenetsk Nats. Region amounted to 1% of the population of the deer 
herds; in 1948, losses reached 2% (Rakhmanin, 1959). In 1951 they 
killed 1708, and scattered 7048 deer which "disappeared without a 
trace". In a decade (1944—1954) these losses amounted to about 75,000 
deer (Semenov, 1954). In 1952 in Arkhangel'sk district, wolves and 
bears killed 1282 horses, 1343 head of large homed livestock, and 
1806 sheep and goats. Of these losses, 64% took place in 7 southern 
regions of the district where wolves were especially numerous. Even 
greater damage to reindeer breeding was caused by wolves. Mortality 
of 3—7% of the domestic reindeer population was from wolves (Korsh, 
1936). Losses of livestock due to wolves in Azerbaidzhan comprised 
20—25% of the total losses in livestock breeding after the war 
(Vereshchagin, 1957). 

These examples show how much the great losses after 1941— 
1945 were caused by the multiplication of wolves. To this must also 
be added the damage which is inflicted by wolves on dog husbandry 
by the destruction of hunting dogs during hunts, sometimes nearly in 
front of the hunter's eyes. Laikas and hounds in particular suffer from 
wolves, which follow their voices. However, wolves also attack watch- 
dogs, falling bn them right in villages, or luring them beyond the 
outskirts. 

Wolves cause great harm to the hunting economy, killing hares, 
foxes, game birds and waterfowl, wild ungulates, as well as muskrats, 
raccoon-dogs and even sables (Koryakov, 1947). Elimination of young 
wild pigs, red deer, roe deer, chamois and Kuban turs by wolves in the 
Caucasian preserve consisted in only 5 months of 34 to 61% (Teplov, 
1938). In Il'mensk preserve, a litter of wolves in 180 days killed about 
100 roe deer, or 15 animals per month on the average (S. Severtsov, 
1941). In the lower Hi river, half of the roe deer and wild pigs bom 
are killed by wolves in autumn. In years with deep snow, wolves may 
kill all of the young (Afanas'ev et al., 1953). In the winter of deep 



266 

snow in 1940/41 in Bashkir preserve, 35 roe deer were found killed by 
wolves, and in the winter of 1942/43, the remains of 8 bull and cow 
marals were met with (not all, of course, were found). A family of 7 
wolves ate a whole bull maral during one night, leaving only shreds of 
the hide (Kirikov, 1952). 

One wolf crossing ice to Barsa-Kel'mes island (Aral Sea) slaugh- 
tered 34 goitered gazelles and 1 saiga from March to June. Moreover, 
beginning in May, it ate the eggs and nestlings of gulls and caught sand 
susliks [Spennophilus fulvus] (Afanas'ev et al., 1953). Wolves cause 
particular harm to wild ungulates in the steppe and desert after calving, 
when they kill the young en masse (Heptner, 1956). 

The intensification of wolf control in the post-war years shaфly 
decreased the "wolf threat". However, it still continues to demand 
serious attention. Even though there is a relatively small number of 
predators, they are dangerous due to their boldness and bloodthirsti- 
ness. Attacking a rich quarry, for example a flock of sheep, the wolves 
kill much more than they need in order to eat. In December 1949, a 
wolf slaughtered 60 geese in one night on a poultry farm in Zhitomirsk 
district (Komeev, 1950). A litter of wolves, especially at the time of 
"hunting training" can kill 60-80 sheep in one flock (Sokolov, 1951). 
191 In the autumn of 1 920, on the headwaters of the Tobol in Kustanaisk 

district, wolves (probably a litter) on a dark night attacked a flock 
spending the night on the steppe, separated it from the shepherd and 
drove off the frightened sheep in the darkness, killing one after the 
other. In the morning, 52 carcasses were found on the steppe, lying in 
a long line in the direction of the flock's flight. Not far away, wolves 
attacked a herd of horses and slaughtered about 10 foals. All the 
animals were killed with a characteristic wolf bite — ^their throats were 
tom (V.G. Heptner). A wolf got into a sheepyard in winter in Redkovichi 
village (Belorussia) and killed 18 sheep in one night (Serzhanin, 1955). 
In June 1941 in the Izhevsk region of Ryazansk district, 2 mature 
wolves attacked a flock of sheep in a corral and in the course of one 
hour slaughtered 96 animals. On the night of 1 January 1950 in the 
Shilovsk region of Ryazansk district, 4 wolves got into a collective 
farm sheepfold and killed 78 sheep (Kozlov, 1955). At the end of June 
1957 in Kopanov village in Ryazansk district, a wolf jumped over a 
wattle fence almost 2 meters high, and burst into a pen near the sheepfoki 
:, where the sheep were driven at night. It killed and wounded 55 sheep, 
of which 5 in all were treatable. There was no watchman and the 
animal worked unhindered. Apparently, it did not carry off anything. 



267 




Fig. 75. Remains of saiga killed by a pair of wolves at a waterhole. Chu river 
valley, Kazakhstan. 3 November 1955. Photograph by A. A. Sludskii. 

After this attack the pen was raised 70 cm and barbed wire stretched. 
After several days, the wolf reappeared and again penetrated the pen 
through a narrow space in the sheepfold wall where the barbed wire 
was not stretched. This time there were watchmen who raised an alarm, 
thereby giving the animal the possibility of a fortunate exit. It wounded 
12 head in all of which 3 could be treated. In this way, the wolf 
destroyed in all 59 head in 2 nights (V.G. Heptner). 

Wolves are the most dangerous vectors of rabies, not only because 
dogs and other animals (and recently, raccoon-dogs) are infected through 
them, but also because the bite of a rabid wolf is, in its consequences, 
about 15 times more dangerous than the rabid dog (Rudnev, 1950). 
Moreover, wolves serve as important vectors of helminths for domestic 
192 animals. The role of wolves in the transmission of the causative agent 
of "sheep gid[diness]" {Multiceps multiceps), and in part echinococ- 
cus, is of great importance. 

Finally, even healthy wolves, although in very rare cases it is true, 
attack humans. More or less verified cases of attacks by healthy (not 
rabid) wolves on people have been registered in Belorussia (Serzhanin, 



268 

1955), in Kirovsk and Irkutsk districts (Timofeev, 1949), in the Kareliya 
(Marvin, 1959), and in the Ukraine (Komeev, 1950). All of them took 
place in winter, the time in which wolves are hungry. 

Thus, in December 1947 in the vicinity of Suna station on the 
Kirov railway, a large man-eating wolf prowled about for one month. 
It usually went out around the village in the morning and evening, 
caught dogs and attacked solitary people, in daytime lying in the bushes 
near the road, not far from the village. In the course of one month, this 
wolf attacked people about 15 times. It bit and scratched 13 people, and 
tore to death two others (a woman and an adolescent). Killed after 
some time this wolf was found to be a large old male (head— body 
length 138 cm) that was extraordinarily emaciated. Except for a bundle 
of woman's hair, nothing was found in its stomach. Apparently, this 
wolf became a man-eater, being unable to catch ungulates. This was 
aided by the fact that during the war, eating of coфses by wolves was 
a frequent occurrence (Marvin, 1959). 

In Ukraine, cases have been described when wolves attacked women 
in Dnepropetrovsk district in the winter of 1923/24 (Barabash-Nikiforov, 
1928); two cases were registered in Stanislavsk district, one in 
Chemigovsk (attack on a shepherd in August), one — ^in Kamenets- 
Podol'sk and one — ^in Poltavsk district, when wolves killed three children 
(Komeev, 1950). 

Control of wolves is carried out in various places by different 
means. Universally distributed are: a) extermination of litters in spring 
and summer in dens, when the pups are not yet grown and do not 
participate in hunting with their parents; b) capture of wolves with 
traps, especially in winter; c) application of poisoned baits. Traps are 
placed in trails ("under the track" or "in the track"), by lures, on scent 
trails, and near the so-called "urination points" (see above). Besides 
poisons which kill the animals (strichnine and others), soporific means 
(for example luminal) have now begun to be used by placing them in 
baits. 

In the forest and forest-steppe zones in the second half of summer, 
autumn, and winter, they are hunted with an enclosure or by driving 
with or without application of flags. Hunting wolves with guns ("when 
howling") is successful. Near baits, and in summer also near the den, 
they stand watch with their guns. In the forest and forest steppe, bor- 
zois and hound dogs are used; in the east they are hunted with golden 
eagles, and sometimes the hunters bring wolves to bay on horseback. 
This method was particularly widespread in Kazakhstan. For this were 



269 

required the best {''baigov")* horses, although wolves do not have 
great endurance when galloping. At full gallop, the animal is killed by 
a blow to the head with a heavy whip (kamchi). 

For this hunting the Ural and Orenburg Kazakhs employed a long 
stick with a root-like knob on the koml*. On such an "ukryuk" [lasso], 
a loop is usually fixed for catching horses from the herd. This method, 
to some extent, takes on a sporting character; however, capturing in this 
way was in the past one of the chief, if not the only method in the 
steppe, and in part, in the forest-steppe habitats of Kazakhstan (V.G. 
Heptner). 

Self-snaring of wolves (loops, frames, cages, pits) are not efficient 
and have local significance; in the Buryatsk ASSR through self-snar- 
ing, 60 wolves were caught in 1946 (ManteifeF and Larin, 1949). In 
Armenia, wolves are caught in winter in the unfrozen irrigation canals 
with quick current and cement walls by building a false bridge (Daf, 
1954). 

The significance of these methods of exterminating wolves is not 
193 equal. According to data derived from a competition on the best means 
of extermination of wolves in 1927, the largest number of predators 
was caught by thorough extermination of litters in dens in summer 
(9.6-5.6 wolves per hunter); in second place was control by poisons 
(7.3-4.7 wolves per hunter); third — driving on horses (5.5 wolves per 
hunter); fourth — capturing with traps (4-3 wolves per hunter); fifth — 
capturing with dogs (3 wolves per hunter); sixth — by shooting from 
ambush near carrion (from 2.7 to 4.7 wolves); and seventh — drives 
using small flags (from 1.4 to 5.2 wolves) (Fedosov, 1939; Danilov, 
1945). 

In 1946, 263 wolf cubs were destroyed in dens in the steppe 
regions of Ryazansk district alone, in Voronezh — ^265, and 637 cubs in 
Astrakhansk. One hunter in Kirovsk district in the summer of 1950 
exterminated 38 wolves by shooting them "when howling"; another got 
17 wolves in this manner in 1949 and in 1950 — 36 wolves. With a 
golden eagle, one hunter caught in 1938/39, 154 foxes and 9 wolves, 
and another, in 1952/1953, caught 124 foxes and 26 wolves. 

During the last few years, in open landscapes (tundra, forest-tun- 
dra, forest-steppe, steppe, desert) and even in the forest zone, the method 
of exteiTTiination from aiфlanes and airsled is found widely applied. In 
the winter of 1945/46 in Kuibyshevsk district, one pilot cornered and 

*Turkic (?) word — Sci. Ed. 



270 




Z5 27 29 31 33 35 37 39 U1 43 «^ ^7 «^ 5f 53 55 
1930 mo /950 

Fig. 76. Graph of tanned wolf hides as an indication of change in wolf numbers 
in the Caucasus by year. Continuous line— Cis-Caucasus; broken line — ^Trans- 
Caucasus (from Vereshchagin, 1959). 

destroyed 180 wolves. Two pilots in the winter of 1947/48 killed 264 
wolves from a PO-2 а1ф1апе in Tatariya and Chuvashiya (Popov and 
Lukin, 1949); in Kalininsk district, in the same winter, 200 wolves 
were caught from a PO-2 а1ф1апе (Kozlov, 1955). In March 1949, in 
Volgogradsk district, 38 wolves were killed from two aiфlanes in 7 
days. In the Nenetsk Nats. Region, up to 1955, the average yearly 
number of wolves captured by all methods was 57 head (including 
cubs), a fact that restrained the growth in the number of wolves but did 
not decrease it. After employment of а1ф1апе8, the number of captured 
wolves grew 8Ьаф1у and the number of wolves and damage from them 
were reduced. Thus in 1955, 92 predators were killed by aiфlanes 
while all land-based hunters caught only 72 wolves; in 1956, 72 wolves 
were taken from aiфlanes, and 48 by land-based hunters (Makridin, 
1959). 

The wolves are very successfully driven and killed by airsled. It is 
applied in combination with an aiфlane (for reconnaissance) and airsled 
in radio contact sent by orders from the а1ф1апе for destruction of the 
wolf packs thus found (N.N). 



271 

194 Genus of Arctic Foxes 

Genus Alopex Каир, 1829 

1829. Alopex. Каир, Skizz d. europ. Thierwelt, I, p. 83, 85. Canis 

lagopus Linnaeus. 
1868. Leucocyon. Gray. Proc. Zool. Soc. London, p. 521. Canis lagopus 
Linnaeus (V.H.). 

Forms of moderate size. 

Skull fairly compact with relatively narrowly placed zygomatic 
arches, quite massive and moderately high (height of the occipital re- 
gion comprises about 1/3 the condylobasal length). Sculpturing of skull, 
particularly sagittal crest, weakly developed, but greater than in g[enus] 
Vulpes. Facial portions moderately elongated (relatively shorter than in 
foxes of g[enus] Vulpes), quite massive but weaker than in the repre- 
sentatives of g[enus] Canis. Distance from line connecting the ends of 
supraorbital processes to anterior end of premaxillae longer than that 
to upper edge of occipital crest. Interorbital (frontal) area convex, 
somewhat elevated in area of frontal sinuses and, on the whole, notice- 
ably and quite abruptly elevated above line of upper rostral profile. 
Supraorbital processes quite weak, flat or even slightly concave from 
above with tapering tip which is directed slightly downwards. Posterior 
edge of tooth row extends to level of anterior border of interpterygoidal 
notch. Deep indentation and protuberance on lower edge of lower jaw 
in front of angular process absent. 

3 14 2 
Dental formula I—; C-; P— ; M— ;= 42. Teeth comparatively 
3 14 3 F J' 

moderate in size and massiveness, camassial well developed; length of 
upper camassial and cheek teeth goes into palatal length about 2% 
times. Canines quite massive, markedly curved and of moderate length 
(when the jaw is closed, tips of upper canines extend to the middle depth 
of mandible, while tips of lower canines reach alveoli of upper). Tips 
of upper canines directed straight downward; no distinct longitudinal 
cutting ridge on posterior side of canine and lateral surface of canine 
smooth. Incisors have no supplemental cusps, or they are very weakly 
developed. 

Body notably elongated, tail quite long — about half the body length, 
ending considerably below the tarsal joint, and in standing animals reaching 
the ground. Head elongated. Ears long with rounded tip. On the soles of the 



272 

hind feet there are only 4 digital pads — behind them there is no large fifth 
pad. 

The fiir is very dense, luxuriant, long and soft. Winter fiir is 
monotone and dimoфhic — ^white, or more rarely grayish-brown; sum- 
mer ftir is dark in color with light flecks. There are two molts per year. 

Supracaudal* gland not developed. From 5 to 8 pairs of teats (A.F. 
Shirkova)'. Evident sexual dimoфhism is absent; age is well marked, 
and seasonal dimoфhism is very shaф, not only in the degree of 
density and length of fur but also in color. 

Forms of the genus possess a quite typical foxy appearance. 

They are inhabitants of the tundra and the Arctic islands. They 
accomplish regular seasonal migrations. They are monogamous, uniting 
together in the reproductive season. Up to 18 offspring (perhaps up to 
20-22 — the maximal number in the family and order), but usually less. 
The young are bom only in a burrow. They are predators, feeding on 
196 birds and small mammals (rodents), flotsam, and carrion. Apart from 
the breeding period, they pursue a solitary way of life. 

The range of the genus is remarkable in that it consists of two 
parts. One of these represents the region of reproduction and is occu- 
pied by the animals in summer time; the other is an area of transience 
and wintering, which they occupy in winter. The region of reproduction 
occupies the whole zone of tundra and the Arctic islands in the Old as 
well as the New World. The region of wintering and winter migrations 
extends on the north to the ice of the Arctic Ocean, and on the south 
to the region of stunted trees [krummholz] and the northern part of the 
forest zone^. 

Moфhologically, the genus is somewhat less specialized than Vulpes. 
Physiologically, as typical arctic [forms], this group is very shaфly 
specialized. The genus stands as a quite well-defined intermediate group 
between the genera Canis (wolves) and Vulpes (foxes). In some re- 
spects, it is closer to Vulpes and has been considered only a subgenus 
of the latter, although it has enough characters that ally it to the genus 
Canis also. More than anything, on the basis of the "intermediate" 

*"Violaceous" in Russian original — Sci. Ed. 

'Reference to 6 (3 pairs) of mammae in the arctic fox, widespread in foreign 
literature, is incorrect. Such a number of mammae is not possible in an animal which 
gives birth to more than 10-12 young. 

'Mammals migrating in a definite season from a more or less considerable part of 
their breeding range are very few. Cases in which the species leaves almost all its range 
are exceptional for the terrestrial mammals if some bats are not considered. Such type 
of migration is quite analogous to the migration of some birds. 



274 

characteristics of the genus Alopex, one may unite all three genera into 
one and consider them as subgenera. It has been assumed that Alopex 
is a subgenus of this broader genus Canis, which includes other groups 
besides the three already named (see above, characteristics of the 
family and of the genus Canis). At present, however, the majority of 
the authors consider the group Alopex as an independent genus. Until 
there is a more detailed analysis of the family system as a whole, this 
point of view is accepted here. 

The genus is known from the Pleistocene (Europe), but it was, 
apparently, established in the upper Pliocene. 

During the period of narrow inteфretation of the species, some 
species were accepted into the genus (some authors considered up to 5 
of them). It is now certain that the genus comprises only one species 
which constitutes about 3.5% of the species of the family. 

In the fauna of the USSR, there is one species, Alopex lagopus 
Linnaeus, 1758; constituting 0.3% of the total species number of the 
fauna. 

Its distribution in the USSR covers the tundra and northern islands 
from the Kola peninsula to Cape Chelyuskin and the Commander 
Islands. 

This genus is one of the most important fur-bearing species in the 
USSR (V.H.). 

ARCTIC FOX [POLAR FOX, PESETS] 

Alopex lagopus Linnaeus, 1758 

1758. Canis lagopus Linnaeus. Syst. Nat., Ed. X, 1, p. 40. Lapland. 
1898. Alopex lagopus spitzbergenensis. Barret-Hamilton et Bonhote. 

Ann. Mag. Nat. Hist., 1, p. 287. Spitsbergen. 
1902. Alopex lagopus beringensis. Merriam. Proc. Zool. Soc. Wash., 

15, p. 171. Bering Island, Commander Islands. 
1912. Vulpes lagopus beringianus^ . Suvorov. Commander Islands and 

fur industry on them. p. 217. Commander Islands. 
1931. Alopex beringensis semenovi. Ognev. Zveri Yost. Evropy i Sev. 

Azii [Animals of Eastern Europe and Northern Asia], vol. 11, 

p. 265. Mednyi Island, Commander Islands (V.H.). 

^This transcription is sometimes wrongly ascribed to Cherskii, 1920 (Ellerman 
and Morrison Scott, 1951). 



275 



197 



Diagnosis 



The only species of the genus. 

Description 

In general appearance, the arctic fox is very similar to the common 
fox {Vidpes vulpes). In its short summer fur, it appears to have relatively 
longer limbs than the fox, with a shorter body. In its luxuriant winter fur, 
it seems to be more squat, although not so extended. The head of the animal 
in its winter fur is fairly rounded, with a somewhat blunt face and very 
widely separated ears which appear slighdy above the fiir; the ear tips are 
rounded. In summer for, the head looks disproportionately large, with large 
and widely separated ears. The tail is -long, about half the body length and, 
in winter fiir is covered with long and luxuriant, extraordinarily dense hairs. 
When the animal is standing, the tail reaches the ground. The soles of the 
feet are covered with dense and long hairs, foot pads are not visible. 

The summer for is. dense, soft and delicate, but short. The general 
tone on the back and neck is smoky-grayish-brown, sometimes with a 
mixture of reddish tones, that on the side is the same but with a lighter 
tone which passes to whitish on the belly. The thighs have the color of 
the sides and the legs, including the feet, are covered with brownish- 
gray for. The dark color of the back in the region of the shoulder blades 
descends like a cape to the limbs and thus forms a cross-shaped figure 




Fig. 78. Arctic foxes, Alopex lagopus L. — ^white (left) and blue. Sketch by 

A.N. Komarov. 



276 

(the summer Arctic fox is called "cross-fox"). The inner parts of the 
limbs are brighter than the outer. The tail above is brownish-gray, and 
the remainder is lighter, mixed with some pale-reddish-brown tints. The 
head is brownish-gray, the dorsal side of the ears is gray-brown and 
their front and inner parts are white. 

The winter fur is extraordinarily long, dense, luxuriant and 
soft. The long and soft guard hairs form a beautiful "veil" of fur, and 
the very firm and dense underfur are characteristic. The color is 
entirely snow white, only sometimes with hardly noticeable creamy 
tints. In addition to this normal white color phase characteristic of the 
great majority of the animals throughout the entire range with the 
exception of some insignificant points, there is also a dark phase, called 
"blue fox". The color of animals of this phase is very variable 
and varies from sandy, gray and light coffee-color to dark ash-gray 
with bluish tint; light and dark chestnut also occur, often with a more 
or less well distinct silveriness (white guard hairs). The whole body is 
198 of one color. The summer fur of arctic foxes of the dark phase does 
not differ, or only slightly differs, from the winter. The summer fur is 
of one tone and the "krestovati/c" ["cross"] pattern* does not occur. 

Sexual differences in color are absent, while age-related ones are 
quite considerable. New-boms are covered with short velvety fur of 
dark-brown color, only very slightly lighter on the sides. With age the 
lightening of color on the sides becomes stronger and gradually form 
the figure in the summer fur described above, and the little one leaves 
the burrow in a "young krestovatik" pattern*. 

For characteristics of the skull and dental system, see above in 
description of the genus. 

The diploid number of chromosomes is 54. 

The size of arctic foxes is subject to quite considerable individual 
variation. Moreover, the average size of individuals of one population 
may vary annually. Extreme and average weights of the animals of one 
and the same population in different years may differ very greatly, 
depending on food reserves which annually change very shaфly, per- 
haps being entirely different. Besides, the nutrient status of animals 
differs at different times in winter — the highest weight is in the begin- 
ning of winter, and by spring it strongly declines. In a good year, 
weight may increase even up to December, and its decrease begins with 
January. There is also geographic variation in size. 

*See above — Sci. Ed. 



277 




Fig. 79. Blue fox, Bering Island, October 1958. Photograph by S.V. Marakov. 



Body length, 46.5-73 cm, tail length 25-52 cm. Weight changes 
from 2 to 8.8 kg (combined data of mainland and island animals, 
including Commander arctic foxes, but without Spitsbergen; see be- 
yond, "Geographic Variation"). In the Lena delta in winter 
(November-March) 1935/36, weight of males (118) was 2400-6800 g, 
an average of 3816 ±* 109 g; that of females (1 14) was 1200-5600 
g, average 3091 ± 107 g; for the whole population the extreme variants 
(232) were 1200-6800 g, average 3453 ± 55 g. In Table 19, changes 
in weight in various months are given (Tavrovskii, 1939). 

Greatest length of skull in males is 127—144.3 mm; in females, 
119—136 mm; condylobasal length of skull in males is 114—138.5; 
in females, 111.0-127.2; zygomatic width in males is 64.7—78 mm; 
in females, 63.2—77.0 mm; length of upper tooth row in males is 

*Omitted in Russian original— Sci. Ed. 



278 






199 



Fig. 80. Skull of arctic fox, Alopex lagopus L. 



279 

53.1-61.3 mm; in females, 50.4-59 mm (skull dimensions include arc- 
tic foxes of the Commander Islands, but exclude Spitsbergen) (V.H.). 

200 Table 19. Weight of foxes in the Lena delta by months (in g|rams|) 



Months 




Males' 






Females 


















Minimum 


Maximum 


Average Minimum Maximum Average 


November 


2400 


5600 


3753 


1200 


4600 


3129 


December 


2700 


5400 


4036 


2220 


5600 


3500 


January 


2500 


6800 


4012 


2000 


4400 


2989 


February 


3000 


4400 


3607 


2400 


3700 


2988 


March 


2450 


3200 


2850 


1900 


3000 


2350 



Systematic Position 

The only species of the genus. 

Geographic Distribution 

The extreme north of Eurasia and North America. The borders of the 
range change considerably seasonally — the region of reproduction oc- 
cupies the tundra and the Arctic islands, in the period of winter migration, 
it covers the northern part of the forest zone and the ice of the southern 
Arctic ocean. The region of irregular winter migrations of individual 
animals extends still farther, both to the north and to the south. 

Geographic Range in the Soviet Union 

This comprises a great part of the range of the species. Within the 
borders of our country is to be found in nearly all the Eurasian region 
of occurrence of the species. It occupies a relatively small part of the 
State territory on its northern frontier. 

The southern boundary of the region of summer occurrence and 
regular reproduction in general follows fairly closely the southern 
boundary of the tundra, but in places, this strict coincidence is some- 
what disturbed. On the Kola peninsula, the above-mentioned boundary 

4x is significant that through January inclusively, the amplitude of weight fluctua- 
tions is very high — the smallest individuals are more than two times lighter than the 
heaviest. In the second half of winter, this contrast is less. Apparently, this takes place, 
to a certain extent, as a result of the decrease in the difference between young and old 
animals at the end of the winter (V.H.). 



280 

proceeds from Varanger Fiord, a little north of Murmansk, to the mouth 
of the Ponoi on the eastern shore of the peninsula. In places, the arctic 
foxes, under the suitable conditions (montane tundra), den somewhat to 
the south of this line. 

On the eastern coast of the White Sea, the border passes a little to 
the north of the mouth of the Mezen', extends across Malozemel'sk 
tundra, mainly its maritime parts, and the northern part of the Timansk 
mountain chain. Crossing the extreme lower Pechora, the border goes 
farther to the east, along the watershed and the upper reaches of the 
Bol'shezemel'sk tundra rivers, i.e. along its middle and southern parts. 
At the Urals, the border extends nearly to the Arctic Circle or a bit 
north of it. On the Urals, the border descends to the south along the 
montane tundras.* 

To the east, the boundary line slightly ascends to the north, reach- 
ing the mouths of Shchuch' and Ob', encircling Salekhard to the north 
and reaches the mouths of Nadym and Pur. Along the divide of the Pur 
and Taz, it descends quite a considerable distance to the south to the 
Arctic Circle. Farther to the east, the border passes northward along the 
right bank of the Taz beyond the Arctic Circle, crosses the Yenisei at 
the level of Noril'sk lake, includes them, and along the mountainous 
places (Putorana mountains?), makes an extension southward, includ- 
ing the sources of the Kureika and Kotue, and apparently Lake Essei, 
not reaching south, however, to the Arctic Circle. 

From the above-mentioned place, the boundary rises quite abruptly 
to the north, crossing the Khatanga in its lower course and proceeding 
201 to the north of 70° N. Lat., crossing the Lena at its lowest reach. 
Farther on, it slightly descends towards the south, crossing at about 70° 
the Yana and Indigirka, and extending south of 70°, reaches the mouth 
of the Kolyma in the area of Nizhne-Kolymsk. From here, this line 
extends to the southeast, reaches the upper Anadyr', and thence abruptly 
turns to the south and even somewhat to the southwest towards 
Gizhiginsk gulf, thus becoming the western boundary for Chukotsko- 
Koryatsk part of the range. The southem border here, i.e. in the direction 
of Kamchatka, passes nearly along 60°, cutting across the very north- 
em part of the peninsula. This is the southernmost known point of 
occurrence (denning) of the arctic fox. 

With respect to information concerning the occurrence of the arctic 
fox in Kamchatka, they are contradictory. According to old data 
(Krasheninnikov, 1 762 and others), the arctic fox was encountered in 

*Not evident on map, Fig. 81 — Sci. Ed. 



281 

the entire peninsula or along its eastern coast to Cape Ukinsk (not far 
from Cape Kamchatka), and along the western coast — to its southern 
extremity (Cape Lopatka) or somewhat to its north. This information 
was inteфreted by some authors as permanent occurrence, and by 
others, as winter migrations. In recent time there appeared a commu- 
nication (Lavrov and Naumov, 1948), that a permanently occupied 
colony of arctic foxes is on the coast of the Sea of Okhotsk in the Ust'- 
Khairyuzov region. All of this information is, for our time, either old 
or doubtful. One may think that the arctic fox is not present now in 
Kamchatka or it is extremely rare. In the past, it is apparent that, on 
the peninsula in the above-mentioned areas, transient animals (regu- 
larly migrating), were caught and farther south there were only rare 
irregular transgressions. 

The northern border of the continental part of the range is every- 
where formed by the ocean. Moreover, arctic foxes den in all the Arctic 
islands from Kolguev and Franz Josef Land to Wrangel' Island, and 
also on Kil'din on the Murmansk coast and on Karagin and Com- 
mander islands in the Bering Sea. Arctic foxes are absent only on 
several very small islands, lying in the open ocean, such as De Long 
Island. However, arctic foxes were found on Uedineniya Island in the 
Kara Sea (V.G. Heptner). 

In spite of some old assertions, aboriginal arctic foxes are absent 
on the Kuril islands. In 1915—1916, blue foxes were brought from the 
Commander Islands to Ushishir (Usisira) Island where they survived 
well and reproduced. In the 20 's and 30 's, arctic foxes were brought 
to the islands of Simushir (Sinsiru), and from the small Kuril islands 
of Yuru (Yurii, Uriri), Kharukaru* and Lis'i (?, V.H.)**. On the Malyi 
[small] Kuril islands at the present time (60 's) foxes are apparently 
absent (Kuznetsov, 1939; V.G. Voronov and A.G. Voronov). 

The southern border of the area of regular migrations — ^the winter 
range — ^in the extreme west passes along the southern coast of the Kola 
peninsula. Within Kareliya, normal migrations are not common, and 
here only occasional long movements are known. The southern border 
of the above-mentioned area between the White Sea and the Urals 
traverses the northern part of the forest zone (V.I. Parovshchikov), i.e. 
somewhat south of the Arctic Circle, although not rarely the arctic fox 
also penetrates far to the south. 

*Kharimkotan?— Sci. Ed. 

**This query by Heptner is not explained, but may refer to the source(s) from 
which the above data are derived. In particular, the context of the sentence suggests that 
foxes were not brought from, but rather, to the small Kuril islands — Sci. Ed. 



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283 

To the east, the boundary of winter habitation crosses the Urals 
and the Ob' somewhat to the south of 65°, reaching the upper Pur and 
Taz, or in the vicinity of their watersheds with Vakh, and crosses the 
Yenisei at Verkline-Imbatsk (below the mouth of the Podkamennaya 
Tungus). Thereafter it goes to the source of the Vilyui and crosses the 
Lena, Yana and Indigirka rivers around the Arctic Circle or a bit more 
to the north, passes over the Kolyma slightly above Verkhne-Kolymsk 
(considerably south of the Arctic Circle) and, still descending south- 
ward, reaches the Penzhinsk Gulf. This district includes the northern 
and western (excepting the extreme south) parts of Kamchatka. 

In general, it may be assumed that winter migrations of the arctic 
fox extend to the northern parts of the forest zone — sometimes quite 
deep into it. In such cases, apparently foxes of the extreme west (Kola 
203 peninsula) roam to the south least of all; there is comparatively little 
shifting of the range in the remaining European part of the [Soviet] 
Union and in Yakutiya, and animals in west and central Siberia move 
southward farthest of all. Evidendy very distant migrations occur in 
places in northeastern Siberia and Kamchatka. 

Migration of separate individuals does not always take place di- 
rectly south- or northward. Many animals also move in other directions. 
In different regions, there may be different predominant directions. 
Thus, arctic foxes from Yamal move mainly to the southwest and 
straight west, going as far as Cheshsk Gulf. Animals during these 
journeys put behind them many hundreds, even more than a thousand, 
kilometers, without even leaving the region of summer distribution of 
the species. This probably explains the fact that in the European part 
of the [Soviet] Union, the border of the winter habitation of the arctic 
fox shifts comparatively little (for more see description of migrations 
in the section "Biology"). 



202 Fig. 81. The range of the arctic fox, Alopex lagopus L., in the USSR 
(reconstructed). The continuous line limits the southern region of summer 
habitation and reproduction (denning), the dashed line — the region of regular 
wintering and permanent winter migrations. Dots indicate places of most distant 
winter transgressions of separate individuals to the south, stars — to similar places 
to the north on the ice of the Arctic Ocean, figures near them give the latitude. 
The question mark in Kamchatka refers to the place (Ust'-Khairyuzovo) regarding 
which there is information about the denning of a small group of animals in an 
isolated spot, triangles — places of Upper Pleistocene finds of arctic fox. V.G. 

Heptner. 



284 

The outlined southern boundary of the region of normal winter 
distribution is entirely tentative. The reason for this is above all an 
insufficiency of accurate materials. Moreover, this line is much less 
shaф in nature than the boundary of the area of denning. Depending 
on various reasons, in one and the same district, the distance of 
migration to the south and the number of animals coming from the 
north, may annually change noticeably. Therefore, the border may be 
situated in the south sometimes far and sometimes near, while at the 
same time, in various parts of the range, these annual variations may 
be expressed in various degrees. Finally, in a series of cases, with the 
presence of data in the literature, it may be difficult to decide whether, 
in a given place, we deal with regular residence of a small number 
of individuals, i.e. with the district of normal wintering, or with a 
place whence separate vagrant individuals migrate comparatively 
frequently, i.e. with a district of long-distance transgressions (in the 
present description, the southern boundary of the normal migration to 
the south is directed, apparently, somewhat more to the south than is 
accepted by some authors, at least for separate parts of the range). 

To the south of the region of regular wintering is situated a wide 
zone of irregular transgression of separate vagrant individuals. They 
often transgress so far to the south, that they, apparently, do not have 
the possibility of returning back, and in the long run they probably 
die. The following most distant places of transgression are known 
(extreme points, from west to east, Fig. 81; for more see the section 
"Biology"): Baltic Sea coast at Ventspils (Vindavy; Van den Brink, 
1958), Daugavpils on the Zapadnaya Dvina (about 56° N. Lat.), up- 
per Velikaya river, flowing into Pskovsk lake, Velikie Luki region, 
northwest of Yaroslavl' (58° N. Lat.), northwest of Kirov at 60° N. 
Lat., the Omutninsk region in the upper Viatka at 58°40' N. Lat., 
northwest of Tobol'sk, approximately at 59° N. Lat., Yalutorovsk on 
the Tobol at 56°40' N. Lat., middle course of the Vasyugan — a tribu- 
tary of the Ob', the mouth of the Parabel' at the Ob', Kolpashovo on 
the Ob' at 58°30' N. Lat., the Chulym river to the north of Achink 
(about 57° N. Lat.), Kansk (west of Krasnoyarsk), Taishet (west of 
Krasnoyarsk at 56° N. Lat.), Bratsk on the Angara, Cheremkhovo on 
the Angara (to the northwest of Irkutsk at approximately 53° N. Lat.); 
one of the southernmost points in Siberia), Bodaibo on the Vitim 
(about 58° N. Lat.), Mukhtui on the Lena (about 60° N. Lat.), 
Olekminsk on the Lena, Yakutsk, mouth of the Tompo river at the 
Aldan, source of the Tompo river (about 64° N. Lat.), Oimyakon 



285 

(about 63° N. Lat.). Tauisk Gulf on the Sea of Okhotsk, southern 
Kamchatka (about 52° N. Lat.), northern Kurils. Migrations to the 
lower Amur and Komsomol' sk-on- Amur — 50° N. Lat. are known 
(points from personal material of A.F. Chirkova). It is possible that 
migrations to the lower Amur may be in part connected with the 
movement of animals along the ice or drifting with floating ice. 
Therefore, in the European part of the country migrations extend to 
56° N. Lat., in the Asiatic — nearly to 50° N. Lat. This difference 
could be explained by the fact that conditions characteristic of arctic 
fox are, in Siberia, distributed farther to the south than in the west; 
in particular, montane tundra extends far to the south. Another reason 
204 is that human settlement in the west is much more dense and the 
animals die earlier, before they attain their natural movement."* 

Winter outposts of arctic fox from the region of reproduction 
occur not only in the south, but also in the north — on the sea ice. 
Movement in this direction represents an entirely normal phenom- 
enon, since in its wandering on the ice, the arctic fox usually follows 
the [polar] bear. It is evident that in southern latitudes and districts 
of the Arctic Ocean, this is a "regular" winter habitation to very 
distant places — ^"irregular long-distance transgressions." However, the 
definition of these districts is still almost impossible. 

As shown by observations in recent years, including both our 
own and also American drifting research stations, arctic foxes are 
more common in the region of the southern sea margin and southern 
parts of the Arctic Ocean, and the farther toward the polar district, the 
rarer they become. Known extreme points of arctic fox finds show 
that they penetrate to the most inaccessible parts of the Arctic Ocean 
farthest from land, and that they probably reach the pole. Very many 
cases of arctic fox observations are known from the ice region rela- 
tively close to the continent (up to 100-150 km). The following are 
the most northerly encounters in the waters in our sector of the 
Arctic: 77°32' N. Lat.— 132°59' E. Long.; 78°19' N. Lat.— 141°33' E. 
Long.; 78°27' N. Lat.— 177° E. Long.^; 78°30' N. Lat.— 176°03' E. 
Long.; 78°42' N. Lat.— 152°33' E. Long.; 80°30' N. Lat.— 159°50' W. 
Long.; 80°5rN. Lat.— 176°00'W. Long.; 85°20' N. Lat.— about 



''Information concerning the capture of an arctic fox in the Tien Shan, which is 
widely found in the literature (for example, Ognev, 1931), and even used for some 
historical formulations is based on a misunderstanding: the skin pertains to an albino 
red fox (Afanas'ev et al., 1953). 

^"Pravda" newspaper on 16 April 1941. 



286 

60° E. Long, (our northernmost point; Chapskii, 1946; Rutilevskii, 
and Uspenskii, 1957). 

All points mentioned are distant from the nearest land by several 
hundred kilometres — in a biological sense, this is more important 
than the latitude. Thus, the place of occurrence at 85°20' N. Lat. 
stands 550 km from Rudolf Island (Franz Josef Land); the point at 
78°27' Lat. lies more than 500 km distant from De Long Island and 
more than 750 km from Wrangel Island; the point at 78°30' lies 750 
km from the New Siberia[n Islands]; that at 80°51'— 900 km from 
land; and the point 80°30' and 159°50' W. Long, found in the "pole 
of inaccessibility", is 1025 km from Point Barrow in Alaska and 1 125 
km from Wrangel Island. In the American sector of the Arctic, the 
arctic fox was noted at 87°, 460 km north of the northern tip of 
Greenland, and at 88° N. Lat., 800 km from Ellesmere Land. 

In the Quaternary period, the arctic fox was distributed much 
farther to the south. Its Pleistocene remains are known in all countries 
of Western and Central Europe, including Ireland, southern France 
(Pyrenees), Switzerland, Czechoslovakia, and Poland. Within our 
boundaries, the remains of arctic foxes are known from the Gomel', 
Bryansk, Lipetsk, Chernigov, Pohava, Dnepropetrovsk regions, and 
even the Crimea. In the east. Pleistocene arctic foxes were found in 
the Chelyabinsk, Krasnoyarsk and Nizhneudinsk regions. Therefore, 
they could be found in the south as far as 44°40' N. Lat. (Crimea) 
and about 55° N. Lat. (Siberia). This demonstrates that contemporary 
transgressions of arctic fox, both in the European part of our country, 
and in the east (Pri-Baikaliya) reach or nearly reach the places of 
their Quaternary occurrence.^ 

It is remarkable that in spite of very ancient and very intensive 
205 exploitation, not considering the comparatively small territory of 
Kamchatka, the range of the arctic fox did not suffer any essential 
changes. 



^Range according to data of Krasheninnikov, 1786; Maak, 1859; Middendorf, 
1887; Pleske, 1887; Ognev, 1926, 1931; Solov'ev, 1927; V.N. Skalon, 1928; Formozov, 
1927; Paramonov, 1929; Yakovlev, 1930; S.P. Naumov, 1931; N.P. Naumov, 1934; 
Tugarinov, Smimov and Ivanov, 1934; Kolyushev, 1936; Heptner. 1937; Sdobnikov, 
1940; Chapskii, 1946; Kuznetsov, 1949; S.P. Naumov and Lavrov, 1948; Stroganov, 
1949; Shvarts, Paviinin and Danilov, 1951; Yanushevich and Blagoveshchenskii, 1952; 
Lable, 1953; N.l. Dement'ev, 1955 and others, and according to unpublished materials 
of V.N. Skalon, V.Ya. Parovshchikov and V.G. Heptner. 

For additional data concerning transgressions, see section on "Biology". 



287 

Geographic Range outside the Soviet Union 

In Europe — the extreme north of Scandinavia and the highest parts of 
the mountains of the peninsula facing the Atlantic Ocean, as a narrow 
strip to the south, nearly to 60° N. Lat. In winter wanderings — ^all of 
Finland, and all Norway and Sweden to its extreme south. Appar- 
ently, they migrate onto the ice of the Baltic Sea. The islands of 
Spitsbergen, Jan Mayan, and Iceland. 

In North America, the southern boundary of the range covers the 
northern parts of Labrador, the Ungava peninsula, and the district 
lying to the east of Hudson Bay. Across the bay and beginning on its 
western shore slightly south of 60° N. Lat., the boundary passes 
westward approximately along this parallel, south around Great Slave 
Lake and along the Mackenzie river to its delta. Farther on, it encom- 
passes the northern and western parts of Alaska, reaching the ocean 
at Cook Inlet. To the north of this line, the range includes the entire 
northern periphery of the continent and all the islands of the Ameri- 
can Arctic and all adjacent districts of Greenland. In the Bering Sea, 
the range includes the Aleutian and Pribilof Islands and Hall, Saint 
Matthew and Saint Lawrence islands. On winter migration, arctic 
foxes reach as far south as the Gulf of Saint Lawrence, and the 
northern parts of Alberta, Saskachewan and Manitoba (i.e. approxi- 
mately 55° N. Lat.). Particularly distant transgressions extend to 50° 
and even 45° N. Lat. (Nova Scotia). (V.H.). 

Geographic Variation 

Compared to the ranges of such species as the wolf or red fox, the 
range of the arctic fox is not greatl Moreover, it is distributed in one 
landscape-geographic zone, and in the various parts of the range 
there are no shaф contrasts in conditions of existence which may 
stimulate geographic variation as an adaptive process. Finally, the 
arctic fox is characterized by seasonal migrations, since some animals 
move for enormous distances, without leaving the region of habita- 
tion and reproduction of the species. This provides a high level of 
panmixia in the species population. Thanks to the continuous cover 
of arctic ice and the wide migrations of animals on the ice, island 



'The exaggerated extent the dimensions of the range of the arctic fox is based 
on the wide use of maps with Mercator projection. 



isolation is pracTically not met with. Within the boundaries of our 
coimtT}-. it exists only in relation to the Commander islands, where a 
separate race w"as formed. In the parts of the range outside the USSR, 
the same conditions and the same tendency towards the formation of 
forms are generally revealed. 

In our part of the range, the following subspecies are distin- 
guished. 

1. Continental arctic fox. A. I. lagopus Linnaeus, 1758. 

Color of winter fur is snowy- white, of summer fur smok\--brown- 
ish-gray (see above): the dark phase is ver.' rare and is usually 
encountered in the propomon 1 : 1000-2000. Changes in this ratio m 
different parts of the range are negligible. Only on Kjl'dm Island on 
Murmansk coast, the percentage of blue reached 30—50. 

SkulP of moderate dimensions, quite broad and high; muzzle 
relatively long (its length constitutes 43-48% of condylobasal length); 
interorbital region wide; infraorbital processes of moderate size; 
postorbital constriction not sharp; braincase quite high and wide — ^its 
height represents 36-43°o. and width — 3A~^l°o of condylobasal length. 
Cranial crests not large, but well defined. Proximal ends of nasal 
bones located at the level of the upper borders of the frontal pro- 
cesses of the maxillar>' bones or somewhat beneath it. rarely higher. 

Size not large; body length 45— ~0 cm. weight to "^ kg. 

Greatest length of skull of males 12"-136 mm. of females 119— 

131 mm; condylobasal length of skull of males 11-1—132 mm. of 

206 females 111-12" mm; zygomatic breadth of males 64. "-"3.". of 

females 63.2—70.3; length of upper tooth row of males 54.1—61.3. of 

females 50.4-59.0. 

Continental part of the range, from the Kola peninsula to Chukotka 
land; all .-Vrctic islands except, perhaps. Franz Josef Land. 

Not recorded outside the borders of the USSR, but perhaps dis- 
tributed m Alaska. 

The Novaya Zemlya arctic fox were, m pan provisionally, some- 
times separated as A. I. spirzbergenesis. This, however, is unfounded. 
for they cannot be differentiated from those of the continent by skull 
dimensions. The abo\e-mentioned form is characterized by ver>" small 
size (condylobasal length of skull 105—120 mm), and infantile skull 
n*pe — the sknill of adult Spitsbergen animals corresponds to that of 

'Characteristics of the skull here and later after Tsalkin (1944). Dimensions of 
skull after Ognev (1931). 



289 



subadult continental ones. That the arctic foxes of the Franz Josef 
archipelago are related, or close to, the Spitsbergen form, is not 
excluded, however, nothing is known of them. 

Craniologically, arctic foxes found in the whole of the above- 
mentioned expanse are identical; however, fur merchants differentiate 
here the following types (Kuznetsov, 1952). 

Novaya Zemlya type. Skins of this type are differentiated by 
especially high quality. They are characterized by large dimensions 
(weight of 100 skins about 30 kg), luxurious silky pelage with tall, 
elastic guard hair and dense underfur, and by the snow-white color. 
Flesh side of skin compact. 

Novaya Zemlya. 

Yenisei type. Skin quality nearly equal to Novaya Zemlya and 
almost the same dimensions (100 hides weigh 28 kg). Pelage only 
slightly less silky. 

Tundra between Taz and Anabara. 

Yakutsk type. Skin dimensions slightly less than the former (weight 
of 100 hides 25.5 kg). Pelage characterized by fluffy, and chiefly, by 
exceptional silkiness; guard hair fine, delicate, less elastic and under- 
fur dense. Color clear-white. Flesh side of skin thin. 

Yakutsk tundra. 

Obdorsk type. Skins of moderate size (weight of 100 hides less 
than 27 kg). Pelage not so fluffy and markedly less silky than that of 
preceding types, with short, especially on the back, elastic guard hair 




Fig. 82. Ratio of for types in tanned skins of white arctic fox in tundras of 

Arkhangel'sk district in the year of movement of arctic fox from the east (of 100% 

at each point) (A.F. Chirkova): 1 — ^Pechora type, characteristic of European 

North; 2 — Obdorsk type, characteristic of Yamal. 



290 

and dense underfur. Color white, but usually with light creamy tone. 
Flesh side of skin compact. 

Region between the Urals and Taz. 

Pechora type. Skin dimensions small (100 hides weigh 26 kg). 
207 Pelage less fluffy, with short guard hair and loose underfur. Color 
white, with barely noticeable bluish tone (color of skimmed milk). 
Flesh side of skin compact. 

Tundras of the European part of [Soviet] Union. 

2. Beringian arctic fox, A. I. beringensis Merriam, 1902. 
Color of winter fur light or dark-brownish, usually with more or 

less well-defined silvery highlights. Dark smoky skins with deep blu- 
ish underfur ("dark blue") are relatively scarce. Fur is extremely 
fluffy, but somewhat coarse, with well-developed guard hair. White 
color phase is absent or found in the form of extremely rare indi- 
vidual cases.^ 

Compared to skull of continental arctic fox, skull narrower in 
region of zygoma and braincase, its height less, supraorbital pro- 
cesses and postorbital constriction more shaфly defined, teeth larger, 
crests more strongly developed. 

Measurements are somewhat larger than continental ones. Body 
length of males 46-67, average 60 cm, females 41-64, average 57 cm; 
weight of males is 3.5-8.8, average 5.5 kg, females 2.5-6.4, average, 
4.4 kg. 

Condylobasal length of skull of males 121-139 mm, females 
116-134 mm. Length of nasal bones in males 41-57 mm, females 
41-50 mm. 

Bering Island. 

Absent outside the borders of USSR. 

3. Mednyi [Copper Island] arctic fox, A. I. semenovi Ognev, 193 1 . 
Color of winter fur of the majority of animals is dark-brownish 

without admixture of white and silvery hairs. Clear-gray ("blue") 
arctic foxes occur in comparatively small numbers. Fur is somewhat 
coarse, with weak, scattered guard hair. White color phase is absent 
or is found in the form of extremely rare individual cases. '° 

"Characteristics of color and ftir, and general dimensions of this and following 
forms mainly after Kuznetsov (1952). 

'"In the past, with suppression of the predominate dark color phase, as on both 
Mednyi island and on Bering island, white arctic foxes were met with more often than 
in the present time. The present situation is the result of the deliberate elimination of 
the white forms over the course of many decades. 



291 

Skull noticeably larger than that of continental arctic fox, narrow 
in zygoma and in region of braincase, its height less, nasal bones 
shorter, supraorbital processes and postorbital constriction more 5Ьаф1у 
defined, teeth stronger. Skull crests, especially lambdoidal and region 
of inteфarietal bone are very robust — ^much more robust than in 
continental animals. 

Dimensions considerably larger than in continental arctic fox, 
and somewhat larger than in Beringian one. Body length of males 61— 
73, average 68 cm, females 60-68, average 64 cm. Weight of males 
4—8.8, average 7.2 kg, females 4-7.2, average 6 kg. 

Greatest length of skull of males 130-144.3, females 122.6-130; 
condylobasal length of skull of males 116-139 mm, females 116—131 
mm; zygomatic width of males 72—78 mm, females 66—77 mm; length 
of nasal bones of males 36-45 mm, females 34-44 mm. 

Mednyi [Copper] Island. 

Absent outside the boundaries of USSR. 

Bering and Mednyi arctic foxes are extremely close to one an- 
other, although the differences between them are quite real, Mednovsk 
are also distinguished from Beringian in averaging shorter nasal bones. 

It is worth mentioning that, at the same time that European island 
races of arctic foxes {spitzbergenensis, fuliginosus) are quite regu- 
larly characterized by their small measurements and by some infantile 
features of skull structure, in Commander (Aleutian and Pribilof) 
arctic foxes, these are not observed. They, on the contrary, are larger 
than continental ones, with more specialized skulls (with features of 
208 hypermoфhosis). Probably, this is explained by the very favorable 
conditions of existence, in the sense of food supply, of arctic foxes 
on the aboveJ-mentioned islands. 

For the part of the range lying outside our boundaries, the follow- 
ing subspecies are usually accepted. 

\) A. I. spitzbergenensis — Barr.-Ham. et Bonh., 1898 — 
Spitzbergen; 2) A. I. fuliginosus Bechst., 1799 — ^Iceland; 3) A. I. 
groenlandicus Bechst., 1799 — Greenland; 4) A. I. ungava Merr., 
1884 — ^Labrador and Ungava; 5) A. I. innuitus Merr., 1902 — Conti- 
nental tundras and islands of America, except the regions occupied 
by the forms ungava and groenlandicus; 6) A. I. pribilofensis Merr., 
1902— Pribilof islands; 7) A. I. halensis Merr., 1900— Hall, Saint 
Lawrence and Saint Matthew islands in the Bering Sea. 

Arctic foxes of the American islands — ^Pribilof and Aleutian, like 
the Commander islands, represent the dark phase. Apparently, not all 



292 

the listed forms are sufficiently well characterized and well grounded . 
(V.H.). 

Biology 

Population. In the tundra zone of the USSR, the arctic fox is the most 
common and abundant carnivore, substituting here for the red fox. 

The number of arctic fox is extremely unstable; it shaфly changes 
by years and by seasons of the year. In autumn and at the beginning 
of winter, in some years, one may encounter tens and even more 
arctic foxes in the course of a day, and in other years, especially in 
spring, only a single individual in the course of a month. 

For orientation, one may assume that the overall arctic fox popu- 
lation in the season before the beginning of harvest amounts, in the 
Soviet sector of the tundra zone, in different years, to from several 
tens of thousands to several hundreds of thousands". Arctic foxes are 
most abundant in the Bol'shezemel'sk, Yamal'sk, Prieniseisk, Prilensk 
and Ust'yansk tundras. 

Arctic fox stocks are not great in the narrow belt of the lokan'go- 
Ponoisk tundra on the Kola peninsula (see Table 20 and beyond). In 
the tundras of northern Arkhangel'sk district, the arctic fox inhabits 
most densely the northeastern Bol'shezemel'sk tundra. In some years, 
these tundras are full of arctic foxes migrating from the east. Arctic 
fox is quite abundant in the north of the Kanin peninsula (Table 21). 
In a year of small numbers of arctic foxes (winter 1951/52) in an area 
of 400 km^ only 3 arctic foxes were taken in a drive; in a year of 
high numbers (winter 1953/54) in several drives on an area of 20 
km^ about 15—30 arctic foxes were taken in December, and in the 
second half of winter about 7—15 arctic foxes. 

In the Malozemel'sk and Bol'shezemel'sk tundras during the 
time of land development work, the general number of burrows was 
revealed (Table 21), the density of which in the northern subzone of 
the tundras increased 3 times. The total number of arctic foxes was 
arrived at (Table 20) by an average "yield", by considering 30% of 
the holes as being occupied and by taking into account two pups for 
each breeding. In these tundras, 7 large "hearths" of reproduction of 

"Boitsov (1937), taking the area of tundra as 3,116,500 km\ considered the 
number of arctic foxes in the 30's to be 100,000 breeding pairs. From this, each pair 
of animals inhabits, on the average, 3 1 km^ or a population density equal to 1 .3 per 
1000 ha. 



293 

arctic foxes were discovered, with the density of burrows ranging 
from 2.9 to 5.6 per 1000 ha in each, and an average of 4.8. The area 
of the hearths comprises about 16% of the entire land of Malozemel'sk 
and Bol'shezemel'sk tundras, and includes about 27% of all discov- 
ered burrows (Skrobov, 1958). In the east of the Bol'shezemel'sk 
tundra, burrows are located at a distance of 1—2 km from each other, 
and rarely at a distance of 400-500 m (A.T. Voilochnikov, Yu.T. 
Nazarenko). At the mouth of the Pechora, in autumn 1944, with 
abundance of arctic foxes close to the multiyear average, hunters 
found in one harvest day 3—10 polar foxes, while in autumn 1954, 
with numbers negligible, 1—2 foxes in 10 days. 

209 In winter 1956/57, on a route of more than 500 km in the region 

of Kara Bay at the extreme northeast of Bol'shezemel'sk tundra, for 
each 20 km of the journey, from 19 tracks in November to 1 track 
in January, averaging 2—6, were found. In winter 1957/58 in a year 
of diminished numbers of arctic foxes was recorded (on a route 
length 1500 km) an average of 0.7 tracks (Chirkova, 1955; Chirkova 
et al., 1959). The fox population in Novaya Zemlya was calculated 
to be several thousand head (Dubrovskii, 1937) (Table 20). 

211 In Yamal, a great number of arctic foxes live in the northern 

subzones of the tundra (Dubrovskii, 1940; Freiberg, 1940; 
Tsetsevinskii, 1940; Chirkova, 1955; Kopein, 1959; N.N. Spitsyn), 
but, apparently, not in the extreme northern edge of the peninsula 
(Table 21), where the tundras exist on a very flat plain. Along the 
Venui-euo, Tiutei, Kharosovoi, Se-yaga, Morade and Tambei rivers, 
burrows of arctic foxes are particularly abundant. At places in the 
middle course of the Tiutei river, burrows are located at a distance 
of only some tens of meters from each other. The region of massive 
denning of arctic foxes in the maritime tundra of the southwestern 
coast of Baidaratsk Bay connects with the best arctic fox lands on the 
Bol'shezemel'sk tundra. Some authors consider that in 1/2 of the 
entire Yamal territory, burrows are few, and probably only in 1/4 of 
the territory are they met with frequently. It is suggested that in 
Yamal, there are about 10 thousand burrows of arctic foxes; this 
constitutes 1/3 compared to the Bol'shezemel'sk and Malozemel'sk 
tundras. Judging by the abundance and catches of arctic foxes, it is 
evident that the number of burrows here is considerably greater (see 
the first footnote to Table 21). 

Already at the end of the last century, Nosilov (1909) who wan- 
dered in Yamal for more than 2 thousand km, could see only 6 arctic 



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296 

foxes, but the Nenety [native people] told him that "arctic fox was 
found not a long time ago and it was encountered on each verst". In 
our time in northern Yamal, in the summer, in a year of average 
"yield" (1949), the number of arctic foxes met with during one day's 
journey by sled* is 1—2, and in winter up to 10 tracks were encoun- 
tered; in the Pri-Ural region in August, up to 10 arctic foxes and in 
winter 2—5 animals. In the Mare-Sale region on the western coast of 
Yamal in autumn, they used to wander in tens, and only after the 
freezing period did their numbers shaфly decline. On the coast of 
Baidaratsk bay, in the middle of August 1947, in a year of abundance, 
on a walk of 80-100 km, more than a hundred arctic foxes were 
counted and in winter, at places for 1 km, there were 1—2 arctic foxes 
and 3—5 tracks. Deep in the tundra, arctic foxes were significantly 
fewer; on a journey extending for 80—100 km, only 10 foxes were 
met with. With an abundance of animals in southern Yamal in the 
winter of 1947/48, the hunters caught from 15 to 35 arctic foxes for 
the season, and in the best conditions, up to 70. In a drive on an area 
of 20—30 km^, 40-60 arctic foxes were taken in December— January 
1955/56 and in February— March, 20—30. In the years when their 
numbers are depressed, on a week's excursion in the tundra, 2—3 
tracks were encountered and only isolated arctic foxes were caught 
in the season; it was only the best hunters in southern Yamal who 
were able to take 10—15 individuals (A.F. Chirkova). 

The total number of arctic foxes in the Yamal region after the 
end of the harvest season for 1955—1958 was 3.1—3.6 thousand head — 
1/4-1/5 of those, that were in the hunting period (12.4-17.6 thousand 
per year). The degree of harvesting was 70-80% of the population 
(Smimov, 1959). However, judging by the number of arctic foxes 
hunted in this region, in which a considerable part of the territory was 
still insufficiently developed for harvest, one may presume that the 
number is considerably greater, especially at the beginning of winter. 

In the northern part of Gydansk peninsula and in the Malyi 
Yamal peninsula, along the Poilovo and Le-Yakha rivers, a large 
concentration of arctic fox burrows is observed. In the territory of the 
latter 21 sections with abundant burrows were revealed by an aero- 
plane. The number of burrows in the Gydansk peninsula and Malyi 
Yamal is assumed to be equal to the number in Bol'shoi Yamal 
(Table 21). But, considering the extent of territory of the Gydansk 

*Russian word is narta, which means both dog- or reindeer sled — Sci. Ed. 



297 

peninsula, the sparse settlement and poorly developed harvest of arctic 
foxes, a significantly greater number of burrows may be expected 
here. 

In the north of Krasnoyarsk territory, arctic fox is comparatively 
densely settled in the tundra of the lower Yenisei, along Pyasina river 
and in part the northeastern Taimyr peninsula (Rutilevskii, 1939; 
Shastin, 1939; A. Romanov, 1941; Chirkova, 1955; Tables 20 and 
21). In the Avamsk tundra, burrows in places are encountered at a 
distance of 10 km from each other. In northern Yakutiya, the western 
regions are richer in arctic foxes than the eastern ones. There are 
many arctic foxes on the New Siberian and Lyakhovsk islands and 
between the Lena and Indigirka [rivers] (Konstantinov, 1921; 
Tugarinov, Smimov and Ivanov, 1934; Freiberg, 1939; A. Romanov, 
1941). 
212 In the Lena-Khatanga territory, the density of burrows declines 

from north to south. In summer, the maritime tundra between the 
Khatanga and Anabara, between Anabara and Olenek north of the 
Pronchishchev range, and between the Olenek and Lena in the region 
of the Chekanovsk range are densely inhabited. More than half of the 
total number of burrows in the Lena-Khatanga territory (6000 bur- 
rows) are located in the northern subzone of the tundra; the shrub 
tundra is the most poorly inhabited. There are many burrows in the 
Lena river delta. Now and then, burrows are met with at a distance 
of 200-250 m to 0.5-1 km from each other, but are usually scattered 
tens of kilometers from each other (A. Romanov, 1941). In unfavor- 
able years, the number of arctic foxes here in winter is not large. In 
Anabarsk and Ust'-Yansk regions, an accumulation of burrows is 
noticed only in some places. In the upper Baranikhi river, near Kolyma, 
F. Vrangel' (1841) found numerous burrows of arctic foxes in the 
past century. 

In the years of low numerical level in the Lena delta, the baited 
traps with an effective radius of about 300 km^ took in from 4—6 to 
30-40 arctic foxes in the winter of 1932 (Freiberg, 1939); the prob- 
able population density — up to 1.3 polar foxes per 1000 ha. When the 
total number of arctic foxes was high in northern Yakutiya in the 
lower Indigirka and Kolyma in the winter of 1946/47, the average 
hunter in one day's harvest encountered up to 13 arctic foxes; the 
ordinary hunter caught about 50, and the best one caught 183, for the 
season. With low numbers in the 1948/49 season on a journey of 20- 
50 km, both in the northwestern and northeastern regions of Yakutiya, 



298 

hunters rarely met with more than 1-4 arctic foxes; of them the average 
caught 10 for the winter, and the best 20-30 animals. Arctic foxes are 
quite numerous on Wrangel' Island (Mineev, 1935; Sergeev, 1936). 

In extreme northeastern Siberia (Magadan district), the average 
density of fox populations in tundra subzones differ by a factor of 10- 
fold (Table 20) (Shustov and Belozorov, 1959). In Bilibinsk region of 
this district in the winter of 1948/1949, in a year of quite large 
numbers of animals, the best hunters took up to 40 animals (Chirkova, 
1955). The arctic fox was scarce in the 30's on the Bering Sea coast 
from Korf Gulf to the mouth of the Apuka river (Koryaksk Nats. 
Region), it was hunted singly and rarely in tens in each region 
(Samorodov, 1939). 

Estimates of arctic foxes on the Commander islands are known 
from the first decades of the present century (Tables 20 and 21). On 
Bering Island, the total number of arctic foxes changed through the 
years from approximately 2 to 4 thousand, and on Mednyi Island was 
in the range of one thousand; the population density on the latter 
island was 2 or more times greater. The considerably great density of 
the arctic fox population on the Commander islands, as compared to 
the mainland, depended at the end of the past century and the begin- 
ning of the current one, on the abundance of marine animal carcasses 
and from the 20's — on the systematic provisioning of the arctic foxes. 

For the historical period in the USSR*, the number of arctic foxes 
changed to the reduced side, chiefly on isolated islands where there 
were numerous blue foxes (Krasheninnikov, 1755; Steller, 1774, 1781; 
Gondatti, 1898; Nosilov, 1909; Formozov, 1929). Mention by 
Krasheninnikov concerning the large quantity of arctic foxes on 
Kamchatka, and Steller' s description (1774) of the innumerable herds 
of these animals in the middle of the 1 8th century, are known; the 
latter killed in one day up to 70 arctic foxes with an axe. The large 
number of arctic foxes on the shores of the Bering Sea in the past 
century and earlier, when arctic foxes in "hundreds ran along the 
shore" depended upon the abundance of marine animals, on the car- 
casses of which, the arctic foxes were feeding. 

There were large numbers of arctic foxes on Novaya Zemlya at 
the end of the 80's and on Kil'din Island in the 70's of the past 
century with the first permanent appearance of humans. One needs to 
assume that the number of arctic foxes on the Kola peninsula and in 

*From the citation dates, this apparently refers instead to Russia — Sci. Ed. 



299 

the northern Far East, also greatly decreased, where the best coastal 
places for polar fox settlement are now densely inhabited by man. 
There are some data about significant decreases in stocks of arctic 
foxes in Sweden (Lost, 1955). 
213 Habitat. Arctic foxes prefer open tundra areas where it also re- 

produces. Only in the years of high numbers and insufficient 
unoccupied habitations do arctic foxes breed among shrubby vegeta- 
tion. The southern border of the denning area coincides with the 
northern limits of distribution of tundra-edge forests. Southward, 
burrows of arctic fox are only found in broad expanses of montane 
tundra. In connection with the climatic warming in the Arctic in the 
course of the last series of decades, a shifting of arctic foxes from the 
southern belt of tundra to more northerly subzones is observed 
(Skrobov, 1960, 1960a). 

The main factors which regulate the distribution of the species in 
the period of sedentary life are the relief of the locality and the food 
conditions. The most suitable places for denning are hilly landscapes 
of dry open watersheds and sources of rivulets and streams; small 



Fig. 83. Tundra along the Khatanga at 72° N. Lat., Taimyr. Core fox habitat in 
summer. Photograph by A.F. Chirkova. 



300 

sandy knolls in open tundra; the upper parts of high terrace slopes of 
creek valleys, the borders of terraces above floodplains, elevated lake 
shores, islands and seashores. Near the southern boundary of the 
range in the summer time, arctic foxes den in treeless rocky slopes 
of mountains, mainly northern exposures, and also in high open water- 
sheds. Burrows here are always situated on elevated sections of tundra. 

Places of denning and feeding usually do not coincide. Denning 
regions change annually. In years with an abundant "yield" of lem- 
mings, a large number of arctic foxes live in the arctic and subarctic 
tundras. With a depression in rodent numbers, arctic foxes remain on 
the seashore, in the southern subzone of tundra and forest-tundra; in 
optimal years, the arctic fox usually does not live in the latter. 

Near the seashore, in connection with better security of food, the 
arctic fox population is more permanent than in other zones. As is 
known, habitats are abruptly changed seasonally throughout the year 
in connection with migrations. Places where small rodents are found 
214 in superabundance serve as feeding biotopes; at any time of year in 
summer, moreover, such places are those where waterfowl occur. In 
winter, arctic foxes also feed along the shores and in areas where 
willow ptarmigan [Lagopus lagopus] are concentrated. 

In the Kola peninsula, breeding places of the arctic fox are re- 
stricted to hilly landscapes at a distance of 40—50 km from the sea. 
The arctic fox constructs the burrow in bearberry-reindeer moss, 
hilly, lacustrine tundra and on alluvial meadows, and forages chiefly 
in willow-grass, sedge, lacustrine, and rocky stand-types, and along 
the seashore. On Kil'din island, it often lives in coastal biotopes 
(Formozov, 1927; Dubrovskii, 1939; Zolotov, 1940). On Novaya 
Zemlya, arctic foxes densely inhabit the low-lying Gusinaya Zemlya 
with its hilly landscape in the southwestern parts of the island; fewer 
arctic foxes occur in montane tundra at Matochkin Shar; in the polar 
part of Sevemyi Island which is covered with an ice shield, arctic 
foxes are found only along the sea coast (Zhitkov and Buturlin, 1901; 
Nosilov, 1909). 

The dispersion of burrows in Kaninsk, Timansk and Malozemel'sk 
tundras is closely connected with sandy-hilly tundra (Shibanov, 1951; 
Maksimov, 1959). In Malozemel'sk and Bol'shezemel'sk tundras, 
93% of the burrows are located on hillocks, knolls, slopes of ravines 
and along the shores of water bodies. Of the burrows 75.5% were 
placed among types of tundra vegetation, of which 22.5% were in 
moss, 18% in lichen, and 16.1%> in shrub formations. 13.8% of the 



301 

burrows were located within marshy tundra, of which the most fre- 
quent type— 9.3%— were found in the vicinity of marshes in 
grassy-tundra, 10.5% were found in shrub tundra, and in meadow 
associations, 0.2% of the burrows (Skrobov, 1958). 

On the Taimyr peninsula, the number of burrows dies away to- 
wards the north, as a function of the transition from the tundra zone 
to the arctic desert zone. On the Chelyuskin peninsula, burrows are 
absent. Burrows are rarely found along the coast of the Kara Sea, are 
relatively frequent along the northern shore of the peninsula and 
along the Nizhnaya Taimyr river. Along the entire western shore of 
the peninsula, the coastal and pre-delta sections, and river and creek 
banks are the areas most densely inhabited by arctic foxes. On the 
eastern coast and on Begicheva Island, their burrows are often situ- 
ated in hilly tundra. In the north of the peninsula, the arctic fox 
reproduces among cotton grass-sedge-moss tundra, where there is a 
rich fauna of rodents and birds, and also along the edge of stony 
deserts, along ravines and in glacial cirques on the montane plateau 
of Byrrang. Burrows are dispersed among dryas patches and grass- 
moss in the arctic semidesert (Koshkin, 1937; L. Lopov, 1939; Shastin, 
1939; Sdobnikov, 1959). 

In the Lena-Khatanga territory, satisfaction of the basic nutrition 
of the arctic foxes is associated with tundra intersected by a dense 
network of small creeks, lakes and laidas*, lake depressions, and the 
seashore, which are the main feeding lands of arctic fox (A. Romanov, 
1941). On Wrangel' island, the arctic fox lives among sandy hillocks 
and feeds along the seashore. 

In the basin of the Anadyr' river, the arctic fox thrives abun- 
dantly in the continental and maritime arctic level tundra, but are also 
abundant in the hilly shrub tundra. There are small numbers of arctic 
fox in montane lichen-shrub tundra and in the forest tundra, and a 
negligible number in the level shrub tundra. In alpine tundra the 
animal is absent. In the montane tundra region of the southeastern 
part of the Koryaksk National Region, the small number of arctic fox 
here are restricted to tundra, the seashore and among shrubs in river 
valleys (Mineev, 1935; Mikhel', 1937; Gassovskii, 1939; Samorodov, 
1939). 

In Magadan district, the density of foxes is highest in the north- 
em maritime subzone; in the southern maritime subzone, its density 

*Openings in the forest-tundra landscape — Gen. Ed. 



302 

is less, and in montane sections of the central interior subzone which 
are far from the sea, its density is lowest. In 1000 ha of different 
types of hunting lands the distribution of arctic foxes is, on the av- 
erage, as follows. In flooded shrubby and meadow tundra — biotopes 

215 favorable for reproduction — ^5 arctic foxes. In complex marshes, in 
the lower courses of large rivers, and along level watersheds and 
maritime plains with abundant lakes — the main biotopes of arctic fox 
in the period of reproduction and raising of young — ^2.25 arctic foxes. 
In the low-lying and transitional marshes and marginal shrub tun- 
dras — 1.3 arctic foxes. In hummocky, cotton grass-sedge, and oiser 
shrub tundra of the northern subzone — 0.75 arctic foxes. In tundra, 
shrubland and marshes of the southern and central subzones of the 
arctic tundra — 0.5-0.25 arctic foxes (Shustov and Belozorov, 1959). 

In Kamchatka, the arctic fox inhabits the zone of sparse stone- 
birch forests. On the Commander islands, it dens in maritime hillocks 
and rocks and, in fewer numbers, lives in the interior of the islands. 
It feeds, mainly, in the laidas, cliffs of bird colonies, and along the 
banks of rivers; in autumn, in tundra and on hills. Throughout the 
year, they frequent human settlements (Suvorov, 1912; Barabash- 
Nikiforov, 1937). 

In winter, the arctic fox is often encountered on ice. During the 
time of migration to the forest zone, they stick to open places — driver 
valleys, watersheds, fields and marshes. 

Food. The arctic fox, like the red fox, is an omnivorous beast, but 
more voracious and less selective; it is poorly provided with food and 
is frequently hungry for long periods. Within the USSR, more than 
125 species of animals and 25 plant species are known to be eaten 
by the arctic fox. 

The nutrition of the arctic fox in continental tundra and on the 
seashore differs shaфly. Already investigators of the past century had 
emphasized the dependence of arctic fox on lemmings (Pakhtusov, 
1842; Krivoshapkin, 1865; Middendorf, 1869; Tret'yakov, 1871; 

216 Silant'ev, 1898). At the present time, numerous investigations have 
shown that the main food of tundra arctic foxes are small rodents, 
which constitute in some seasons and years more than 99.0% of the 
stomach contents and feces; mainly the species represented are Nor- 
way, Ob and hoofed [= collared] lemmings (Lemmus lemmus, L. 
sibiricus, Dicrostonyx torquatus). Gray (genus Microtus) and red 
[backed] (genus Clethrionomys) voles are eaten by arctic fox in smaller 
amounts; these rodents are met with in the arctic fox's diet mainly in 



303 

the southern tundra belt, in forest tundra and in the taiga zone (during 
migration). In places of water vole (Arvicola terrestris) abundance, 
arctic fox feeds also on them; they rarely catch arctic hares which are 
particularly characteristic of the shrubby growths on the 
Bol'shezemel'sk tundra (up to 43% in studies). 

Birds are caught more often in spring and summer (in Yamal up 
to 52% occurrence in analysis). They destroy nests, eating the eggs 
and nestlings; in the second half of summer, they hunt molting geese, 
most frequently the white-fronted goose, and rarely, ducks. In autumn 
and winter, they catch willow and rock ptarmigan [L. lagopus, L. 
mutus], rarely snow buntings [Plectrophenax nivalis] and longspurs 
[Calcarius lapponicus], and still more rarely, birds of prey wintering 
in the tundra. Quite often they eat fish (up to 41% occurrence in the 
food analysis), usually refuse of fish harvest or bait; it sometimes 
catches migrating salmon during spawning time. 




2 1 5 Fig- 84. The shore of Kara Bay (Kara Sea) at the beginning of winter. A place of 
massive migration and harvest of the arctic fox. Photograph by A.F. Shirkova. 



304 

Wild and domestic reindeer are of great importance in the nutri- 
tion of tundra arctic foxes, herds of which arctic foxes usually follow. 
In the diggings [feeding craters] of reindeers, the arctic fox catches 
small rodents; near the reindeer herd, ptarmigan are concentrated; 
fallen reindeer become food for arctic fox (up to 68% in food remains 
in famine years). In winter, arctic foxes eat the feces of reindeer. 
Sometimes, arctic foxes pursue reindeer weakened by a prolonged 
period of ice-covered ground or injured reindeer; they devour lumps 
of snow moistened with the blood of these animals; they lie around 
the dying animals in expectation of their death. Cases are known 
when arctic foxes killed and ate new-bom reindeer fawns, even in the 
presence of their mothers. 

Most of the above-listed foods are eaten by the tundra arctic fox 
only in the absence of lemmings. One must consider them secondary. 

Invertebrates, especially marine, are insufficiently known. In- 
sects occur rarely and in negligible quantities in the food of the arctic 
fox — beetles, bumblebees, wasps, grasshoppers. Extremely rarely, the 
arctic fox eats amphibians (grass frog on the Kanin peninsula). Weasel, 
ermine, shrews, insects and amphibians must be assigned to the class 
of accidental and emergency food, eaten in the years of lemming 
scarcity. 

Cannibalism is developed to a strong degree among arctic foxes. 
In some years, more than 50% of those animals falling into traps, 
were found to be chewed. Cases of eating dogs and red foxes are 
extremely rare'^. 

Arctic foxes inhabiting islands and coasts feed on the flotsam of 
the sea — carcasses of marine animals and birds. They gnaw a way 
through the whole system of a whale's carcass and sometime do not 
come out of it for some days (Nosilov, 1909; Mineev, 1935): they 
gorge themselves with blubber until they vomit. On the littoral, they 
gather cast-up cod, haddock, sea-hen, capelin, arctic cod, navaga 
[Eleginus navaga], smelt and other fishes. There, they also gorge on 
echinoderms, crustaceans, molluscs, sea-cucumbers, ascidians, sponges, 
hydroids and others. In a period of massive consumption of sea ur- 
chins, the teeth of arctic foxes on Mednyi Island are stained violet in 
color. Bird colonies are preferred foraging places for arctic fox. They 
capture black guillemots [Cepphus grylle], murres [Uria sp.], fulmars 
[Fulmarus glacialis], razorbills [Alca torda], gulls and even the 
stronger skuas [Stercorarius], cormorants, goosanders [Mergus] and 

'^0.4-2.0% in analyses of food (Pereleshin. 1943; Skrobov, 1958). 



305 

others. In one feeding, a hungry arctic fox may eat up to 0.5 kg meat, 
fat and fish (Skrobov, 1958). The gastro-intestinal tract of the arctic 
fox fully rids itself of food remains within 24 hours (Pozgnyakov, 
1957). 

217 Arctic foxes feed also on plant food; in the tundra on various 
fruits— crowberry, blueberry, whortleberry, bearberry, cloudberry, 
various species of sedges and grasses, and on the seashore, on 
seaweeds; the significance of the latter is not entirely clear, since it 
is evident that they are not digested at all. Arctic foxes chew on 
russula mushrooms. In the alimentary tracts of 898 arctic foxes were 
found 78 different species and forms of plants taken during feeding 
on other food materials (Pereleshin, 1943). For the arctic fox, as for 
the red fox, ingesting inedible material — discarded reindeer skin, raw 
strips, rags, string, paper, woodchips, crushed pieces of coal, pieces 
of soap, as well as pebbles, sand, glass fragments and others- — is 
characteristic. 

On moving in autumn and winter to the ice of the Arctic Ocean, 
arctic foxes (especially the old ones, for which independent foraging 
is difficult) usually follow polar bears, gathering its food leavings, 
and also remains of the food of ringed seals; they gnaw at the eruc- 
tations of the walrus — a whole pile of indigestible food. Arctic foxes 
return in spring from the ice seriously emaciated. 

The prevalence in the diet of these or other food items depends 
upon the season of the year. Tundra arctic foxes most frequently 
consume lemmings at the end of summer, in autumn and in the first 
half of winter, i.e. at the time of their mass appearance. The most 
stressful time of the year in tundra is the end of the winter and 
beginning of spring. The long duration of ice-cover, often in the 
second half of winter, results in the emaciation and death of reindeer, 
thus providing additional food for arctic foxes; at that time, the ice 
crust which forms renders difficult the excavations of voles and causes 
the death of small rodents. At that time, arctic foxes with broken 
claws are not infrequently encountered. In spring and summertime, 
the possibility of the foxes' feeding at the expense of birds increases. 
In the second half of summer, the molting birds appear, and in au- 
tumn berries ripen. 

The diet of the arctic fox varies in different geographic regions. 
In lokan'go-Ponoisk tundra of the Kola Peninsula (Table 22), at the 

2 1 8 time of rodent reproduction, the arctic fox most frequently feeds on 
Norway lemming (to 96% of all data), they often eat gray and red- 



306^ 





"^ ~-~,^^^ Time of year 
Types of food — ~»,__^ 


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2 1 7 Fig. 85. Change in the composition of arctic fox diet during the course of the year. 
A — mainland arctic fox of Kola peninsula (Zolotov, 1940); В — island arctic fox 
of Kil'din Island, Murman (Formozov, 1929; Zolotov, 1940). 

backed voles, birds and insects. On Kil'din Island in the summer of 
1927, foxes mainly existed at the expense of the shore fauna due to 
the scarcity of rodents (Fig. 85; Formozov, 1929). 

In the northern parts of the Kanin Peninsula (Tables 23 and 24), 
feces collected near the den burrows of arctic foxes contained, almost 
exclusively, mammalian remains. The majority of them belonged to 
collared and Siberian lemmings, and in lesser quantity, voles and 
others. In a year when lemmings were absent, cloudberry was pre- 
dominant among plant foods. In Bol'shezemersk and Malozemel'sk 
tundras (Table 23), the diet of the arctic fox consisted most frequently 
of mouse-like rodents, including Siberian lemming, and when the 
latter was in small numbers, narrow- skulled vole {Microtus gregalis) 
and water vole (at the mouth of the Pechora); moreover, white hare 
and, among birds, chiefly willow ptarmigan (probably the latter, as 
well as hares, represented bait from traps*). Remains of shrews 

*The Russian phrase, "nazhivky iz samolovov", probably refers to bait from a 
dead-fall trap; see Figs. 96, 97 — Sci. Ed. 



307 



(winter, 1957/58, up to 8% occurrences) were found in arctic foxes 
caught near Karsk Bay. Fox stomachs in the winter of 1956/57, when 
lemmings were absent, contained much garbage and household refuse. 
Around burrows of the arctic fox in summer are often found wings 
and bones of willow ptarmigan, and rarely, of ducks, small shore- 
birds and small passerine birds and among mammals, the white hare 
(Skrobov, 1958). 

The nutritional condition of arctic foxes in northeastern 
Bol'shezemel'sk tundra depended on the nutritional conditions of the 
year, as well as on sex, age, and physiological condition. In winter 
1956/57, distinguished by low numbers of small rodents and large 
numbers of arctic foxes, males were in better condition in autumn, 
while during winter they lost weight. Females, especially those with 



Table 22. Diet of arctic fox on Kola Peninsula (in % of occurrences in number 
of investigated objects) 



"^^^.....^^^^^ Place of collection, 


lokan'go 


-Ponoisk tundra 


Kil'din 


«...«season, year ~ 




1935 




Island, 

1927, 
Summer^*^ 


Food ^"^-'---^..^^^^^ 


Summer' *• 


Winter"* 


August" 


Small rodents 






80 


20 


Lemmings 

Gray voles [Microtus] 

Common and Northern red-backed 


96 
46 
42 


92 
15 
13 






vole [Clethrionomys] 
Arctic hares [Lepus timidus] 
Shrews [Sorex] 




1.3 
1.3 






Carrion (reindeer) 
Birds 


42 


9 
29 


18 
16 


40 


Fish 






10 




Insects 


29 






Less than 5 


Marine animals of the littoral zone 








2522 


Crustaceans 






2 




Molluscs 






2 




Plants 


,721 




4 


Less than 5 


Refuse 




il 






Number of small rodents 






Very 
low 


Low 



"*L.N. Borodin. 
'"Zolotov, 1940. 
^°Formozov, 1 929. 
^'Berries. 
^^Mostly sea urchins. 



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53 



310 

219 litters, were more emaciated in autumn, but in winter, they fattened 
up; only a few specimens were characterized by severe emaciation — 
puфlish color of muscles and great projection of bones. At the same 
time, arctic foxes which were, apparently, feeding on the coast, were 
noted to have large fat deposits — ^subcutaneous fat on the sides up to 
2—3 cm, and in the form of layers in the muscles. From one male, up 
to 1 kg of fat was removed, which represented 1/4 of its carcass 
weight. In the following winter 1957/58, distinguished by abundance 
of lemmings and low number of arctic foxes, the young females were 
very well nourished, but those which reproduced became of lower 
than average nutritive condition (Chirkova et al., 1959). 

Foods of the arctic fox on the islands of Novaya Zemlya are 
varied. The animals move back and forth everyday from the tundra 
in which they catch lemmings, to the coast where they feed on marine 
flotsam. In rivers, they catch chars and live around the carcasses of 
marine animals for long periods in snow burrows. Hunting in bird 
colonies, they destroy completely the accessible nests. In Franz Josef 
Land, the arctic fox feeds mainly at the expense of bird colonies 
(Nosilov, 1909; Esipov, 1935). 

220 Yamal arctic foxes, far from the coast (Table 25), fed exclusively 
on terrestrial mammals (in 1939—1942 up to 99%— 100% of occur- 
rences in diet; Dunaeva and Osmolovskaya, 1948), and sometimes 
uninterruptedly on mouse-like rodents. In summer 1933, in the Tambei 
river basin, the latter formed 99.6% of occurrences in feces at the 
burrows (Tsetsevinskii, 1940). They were mainly Siberian lemming 
(up to 79.5% of occurrences from total sample amount and up to 
92.5% of all rodents in northern Yamal), and fewer collared lem- 
mings, and also gray and red-backed voles. The importance of 
lemmings, especially the Siberian, increases as one moves northward 
on the peninsula. At places in the floodplain of the Ob' river and Ob' 
Bay, water voles occupy some place in the diet of the arctic fox. The 
portion of birds in the diet of tundra foxes with lemmings abundant 
is not great even in summer, but in individual years in the taiga belt 
of the Yamala-Nenetsk National Region, it increases greatly (to 50%). 
Here, the arctic fox gorges on fish, mainly the sorts of refuse from 
places of fishing harvest. In a year of abundant berry crop and few 
lemmings, plant food is of great importance and also carrion. 

Great fluctuations in the dietary composition is observed in the 
tundra arctic fox, depending on annual abundance of lemmings (for 
example, winter 1932/33 and 1933/34— Table 25: Tsetsevinskii, 1940). 



311 

In winter 1938/39, in southern Yamal (Korzinkina, 1946), the occur- 
rence of rodent remains in the arctic fox diet was observed to grow 
monthly in connection with subnival lemming reproduction. Bird re- 
mains also declined. With an abundance of rodents in autumn 1939 
and their gradual disappearance in winter 1939/40, the occurrence of 
rodent remains in the stomach contents diminished monthly; at the 
same time, the arctic fox fed more often on reindeer carrion and 
caught birds. 

On the Taimyr peninsula, the principal food of the arctic fox also 
consists of small rodents. In August 1931 on the right bank of the 
lower Khatanga river, among the bones of mouse-like rodents col- 
lected near an old den burrow, 85% of the skull remains consisted of 

222 collared lemming, and 15% — Siberian (A.F. Chirkova). In winter 
1932/33 in the neighborhood of Khatanga village and a series of 
eastern stations* (Table 26; Skalon, 1936) with a lemming die-off, 
small rodents occurred in 68% of the dissected stomachs of arctic 
foxes; predominantly voles — M. middendorjji and M. hyperboreus 
and the Siberian lemming. The remains of reindeer carrion, fish mainly 
from baits, and also willow ptarmigan and some other birds consti- 
tuted the three other types of food; each of them was found in nearly 
1/3 of the stomachs. The considerable mixture of food and inedible 
human household refuse was also an indication of the lack of small 
rodents in that winter. 

On the southwestern coast of Taimyr and on Begichev island, the 
principal food of the arctic fox was also recognized as lemmings, 
especially the Siberian. On the Chelyuskin peninsula, in a lemming 

223 year, the remains of collared lemmings — the only one occurring here — 
constituted the principal content of tundra arctic fox feces. In years 
when rodents disappear, the diet of the arctic fox mainly consisted of 
marine flotsam which is not present here each year owing to the 
different times of ice formation, and of the leavings of the polar bear. 
Arctic foxes fed in autumn 1932 on marine flotsam. The foxes also 
dug this food out from under the snow in winter, living in a series of 
snow burrows. In winter time, arctic foxes here frequently do not get 
enough to eat, and in connection with this cannibalism develops among 
them. In the famine winter of 1944/45, arctic foxes dug up the lem- 
ming carcasses stored from the preceding year, when these rodents 



*In the original Russian text, this word is stankoy (gen. pi.; machine-tool); 
apparently a misprint for stantsov — ^Sci. Ed. 



312 

220 Table 24. Mammalian species in food remains of the arctic fox on Kanin 
Peninsula in summer at burrows (in % of total number of occurrences 
in feces; Shibanov, 1951) 

Quantity 554 604 



Food ^"^--^^^ Year 1946 1947 

Collared lemming [Dicrostonyx torquatus] 56.8 48.9 

Siberian lemming [Lemmiis sibiricus] 21.1 8.4 

Wood lemming [Myopus schisticeps] 0.5 0.2 

Lemmings, not accurately identified 0.4 — 

Root vole [Microtus oeconomus] 3.3 4.0 

Northern red-backed vole [Clethrionomys rutilus] — 0.3 

Voles, not accurately identified 10.4 30.4 

Mouse-like rodents, not accurately determined 5.7 6.6 

Shrews [Sorex] — 0.2 

Arctic fox 1.8 1.0 



100.0 100.0 



were abundant, and died after that (К1ф1сНп1коу, 1937; Koshkin, 
1937; Rutilevskii, 1939; Sdobnikov, 1958). 

In Lena-Khatanga territory (Table 26) during a зЬаф reduction 
in numbers of mouse-like rodents, mainly collared and Siberian lem- 
mings were found in the stomachs of half the arctic foxes in winter 
1926/27. The foxes took birds and fish more often from lures (A. 
Romanov, 1941). In the lower Lena, the arctic fox also preys on 
northern pika — Ochotona hyperborea (Kapitonov, 1961). 

In the eastern regions of northern Yakutiya (Table 26), the years 
in which the investigation of arctic fox diet fell apparently coincided 
with few rodents (Krasovskii, 1939). In the delta of Lena, the arctic 
fox frequently fed on fish and birds. 

The existence of arctic foxes on the Commander islands is asso- 
ciated completely with the seashore. Their diet is composed (Table 
27) of the carcasses of marine mammals, especially fur seals 
224 [Callorhinus ursinus] (up to 56% in the analysis), birds, echinoderms, 
mostly sea urchins, and other littoral invertebrates, fresh fish, and 
dried sahnon from foodstores, crustaceans, and molluscs. Sometimes, 
arctic foxes capture young fiir seals from their mothers, or drive away 
ringed seals from their breathing holes into the ice and then attack 
them. Cannibalism is not observed on the Commander islands. Among 
plant food, the arctic fox eats berries in the tundra, laminaria kelp. 









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317 



Table 27. Arctic fox diet on Commander islands (in % occurrence to number 
of investigated objects to total of all occurrences) 



\ Place of collec- 

^v tion, number of 

N. objects, season. 


Bering 


Island 


Mednyi 


Island 


194 


534 


250 


296 


>y^ year 


stomachs. 


stomachs, to 


stomachs, to 


stomachs, to 


\^ 


to the 


number of 


total of all 


number of 


>v 


total of all 


investigated 


occurrences. 


investigated 


Food \^ 


occurrences''^ 


objects, 
1929/30« 


1924/25^' 


objects, 
1930/32^5 


Small rodents 


1.1 


0.4 






Marine animals 




0.2 


56 


6.8 


Birds 


2.1 


2.2 


4 


39.5 


Fish 


38.4 


10.3 

63.5^^ 




14.9 


Littoral marine 


32.4 


15.7 


2049 


30. n-* 


organisms 








16.2 
others 


Crustaceans 




3.4 




25.0^" 


Molluscs 


2.5^^ 


1.1 


1 


23.0 


Supplemental 

foods, lures, baits. 


2.7 




8 


13.5 


household refuse 










Plants 


10.3 


8.0 
19.1^** 


24« 


14.5 


Inedible (rags, leather, 




0.6 






nets, minerals, etc.) 










Empty stomachs 


10.5 


5.0 


8 




Total in % 


100 




100 





«Cherskii, 1919-1920. 

«Lavrov, 1932. 

''^Freiberg, 1929. 

''^Barabash-Nikiforov, 1937, 1939 (diet of tundra and not bait-fed arctic fox). 

''^Octopus, dvuustika* 

"'Dried fish — salmon 

'"^Sea cabbage. 

"^Mostly sea urchins. 

^"Insects— 0.7%. 



'Lit., trematode fluke; meaning unclear in this context — Sci. Ed. 



318 

rockweeds [Fucus] and other seaweeds of the httoral (Cherskii, 1919— 
1920; Freiberg, 1926; Lavrov, 1932; Barabash-Nikiforov, 1937a, 
1939). 

There are age and sexual variations in the nutrition of arctic 
foxes. Juveniles more often feed on lemmings. Among the Khatanga 
arctic foxes (Skalon, 1936), the lemmings are more often eaten by 
females than by males. Males caught mainly gray voles and also fed 
more on reindeer carrion or on baits of birds and fish and as a whole 
they did not show selectiveness in food. East of the Lena (Krasovskii, 
1939), lemmings were also mostly found in the food of females. On 
Belyi island, differences in the diets of males and females were not 
observed. 

Home range. A definite home range is characteristic of the arctic 
fox only in spring and summer, during the time of reproduction and 
growth of the young. On the Commander islands, at the time of den 
establishment and then the emergence and raising of the young, the 
arctic fox occupies a permanent territory sometimes demarcated be- 
tween natural boundaries in the form of streams, knolls and other 
particular relief features. In the event that a "foreign" arctic fox 
intrudes, it is repelled by the parents; in this case, even the weakest 
female can drive away the stronger newcomer. However, the area 
does offer complete security to the whole family, as the parents go 
for food to the seashore or to other regions (Cherskii, 1919-1920; 
Boitsov, 1937; Barabash-Nikiforov, 1939). Thus, the home range is 
as much a breeding as it is a feeding territory. 

The dimensions of the selected territory vary according to the 
locality and abundance of food. In rugged terrain, the areas are smaller 
than on the plains. In the delta of Lena, the area inhabited by a family 
in the summer is from 5 to 30 km^. In the northern part of the 
Kharaulakh range, dispersal of the arctic fox from the burrow in June 
1927 did not exceed 5 km; the same was also observed on the plains 
in the lower Anabara river in 1934. Both years were characterized by 
low numbers of lemmings. On the Taimyr, in the region of Ozhidanie 
bay, in the year of a good "yield" of arctic foxes (1947), the area of 
their breeding territory comprised not less than 16-25 km^: 3 burrows 
with litters were found at a distance of 4—5 km from each other (A. 
Romanov, 1941; Sdobnikov, 1958). Breeding territory on the Kanin, 
defined as the distance between burrows, was equal to 2—14 km^ 
sometimes more in North Yamal — up to 25-30 km^ and in the Lena- 
Khatanga territory — ^from 20 km^ and more. On the Bol'shezemel'sk 



319 

and Malozemel'sk tundras, in 7 centers of arctic fox concentration in 
an enormous territory of 16,500 km^ in a comprehensive tally each 
2 km^ contained only one burrow on the average (Skrobov, 1958). 
With food abundance and dependability, litters of these animals are 
sometimes placed in immediate proximity to each other, as was ob- 
served on the Yamal, Kanin, and in Belyi and Mednyi islands 
(lokhel'son, 1898; Freiberg, 1926, 1939; Tyulin, 1938; Tsetsevinskii, 
1940; A. Romanov, 1941; Shibanov, 1951; Sdobnikov, 1958; Skrobov, 
1958; N.N. Spitsyn). 

As soon as the young emerge on the surface, arctic foxes often 
roam from one burrow to another, especially if they are disturbed. On 
the Commanders, parents accompany the young to the seashore to 
temporary shelters (Cherskii, 1919-1920; Barabash-Nikiforov, 1937a; 
Maksimov, 1959). 

After the young arctic foxes establish themselves at the end of 
summer and beginning of autumn, the boundaries of the littering sites 
cease to exist. At this time, the arctic fox commences a nomadic life. 
The distribution of arctic foxes after that depends on the abundance 
of food in autumn— winter period. In the years with bad supply of 
225 food, the arctic fox moves for some hundreds, thousands and even 
more kilometers. In years of lemming abundance in continental tun- 
dras, arctic foxes in summer restrict themselves to region of summer 
denning during winter also, and their individual ranges at time are not 
large. In the Lena delta in a year of lemming mortality, arctic foxes 
which remained to winter in the tundra moved for a distance of up 
to 10 km in the direction of baited traps (Freiberg, 1939; A. Romanov, 
1941). 

In the Bering and Solovetski islands, constancy of occupation 
from year to year of different places by definite groups of arctic foxes 
distinguished by their fur color and behavior was noted. 

Burrows and shelters. The choice of a place for construction of 
the burrow, and then its use for many years, depend on many factors. 
Chief are the dryness of the soil and the absence of proximate subsoil 
water and permafrost, slope exposure, the lookout possibilities of the 
place, and food supply in summer time. Arctic foxes' burrows are 
usually located on elevated dry interfluves, on small hills, precipices, 
etc. 

In the mountains or in outcrops of bedrock, the arctic fox digs a 
burrow in a fine-grained layer, and if it is absent, they settle in rock 
clefts and between plates of montane bedrock. On Kil'din Island, 



320 

burrows and dens of arctic foxes are concentrated in a coastal belt 1- 
2 km in width or narrower; here arctic foxes live at various altitudes, 
from some tens of meters above the water to the extreme tops of the 
plateau. Burrows, dug in sand, were abandoned because of frequent 
human visitation and their destruction, and the foxes burrowed within 
rocky areas under plates of bedrock (Formozov, 1929). 

The arctic fox also uses the burrows of other animals, for ex- 
ample, the marmot [Marmota camtschatica] (Kharaulakh range; V.I. 
Kapitonov). In Magadan district, it burrows in places with old ground 
squirrel colonies with softer, drier soil (Shustov, Belozorov, 1959). In 
seabird colonies on Novaya Zemlya, arctic foxes dig burrows in the 
guano (Nosilov, 1909). 

In tundras where wind force is great, sheltering the burrow from 
it is important. In the northeastern Bol'shezemel'sk tundra, burrows 
are more often placed at a height of 2/3 up the slope, not exceeding 
12 m above water level of the nearest water body. In such a place, 
wind force near the burrows is reduced by 32% of its force on the 
slope crest (Danilov, 1958). In the zones of open tundras in the 
northern belt, the arctic fox prefers south slopes, without deep snowy 
faces, which thaw out earlier in spring, and which are protected from 
northerly wind in summer time. In the Bol'shezemel'sk and 
Malozemel'sk tundras, of the 483 burrows inspected, 26.5% were 
located on the southern side, 17.8% on the eastern, 14.3% on the 
southwestern and 11.3% on the northern side. To the north and near 
to the coast, the number of burrows on south slopes increases. Thus, 
in maritime tundra of the Taimyr peninsula, almost all burrows are 
located on slopes of southern exposure, while in the upper reaches of 
the Khantaiki, Kureiki and Kotuya rivers below 67°30' and 68° N. 
Lat., arctic fox burrows are restricted to the northern slopes (N. 
Naumov, 1934; Sdobnikov, 1958; Skrobov, 1958). 

Burrows are constructed in the vicinity of water bodies, usually 
within 100-500 m. Paths lead to the water. In the southern zone, 
burrows are located near growths of shrubs in which young arctic 
foxes hide from enemies and bad weather, and where they can also 
find food in the form of ptarmigan and small rodents. 

The arctic fox constructs permanent dwellings, usually in sandy 
and sandy loam soils, more rarely in sandy-gravel mixed with pebbles 
and still more rarely in clay soil. In summer, the soil temperature in 
burrows is some degrees higher than outside. However, Middendorf 
(1869) encountered in the Taimyr peninsula burrows in which even 



321 

in the middle of summer, icicles hung like stalactites, with arctic fox 
wool frozen to them. 

The arctic fox digs burrows of no great depth, from 0.5 to I m, 
rarely more. On Belyi Island, in the Taimyr and in the Lena-Khatanga 
territory, they are distributed 1 0—1 00 cm above the permafrost (Tyulin, 
226 1938; A. Romanov, 1941; Sdobnikov, 1958). In places with burrows, 
as compared to neighboring areas, the level of permafrost is lower. 
At Cape Osten-Saken on the northwestern shore of the Taimyr, in a 
place where burrows are found, the permafrost level is restricted to 
a depth of 65 cm (bottom of burrow to 55 cm), while to the side of 
the burrow, [frost] depth is 25 cm. 

Complexity of the burrow increases through use from year to 
year. As a result of quite rapid breakdown of burrows, arctic foxes 
build new passages and the center of the lair gradually shifts. The 
breakdown proceeds as a result of settling of friable soils through 
constant digging; this is also favored by exposure to the air and 
trampling of the burrow by reindeer herds attracted thence by succu- 
lent vegetation, and also by the digging of humans. The more favorable 
place the burrow is located, the more often it is inhabited by a breed- 
ing pair. For that, arctic foxes each time undertake a cleaning of the 
holes and exit passages. Usually, arctic foxes renew 5—10 passages 2— 
8 m in length. In Bol'shezemel'sk and Malozemel'sk tundras, out of 
483 burrows inspected, temporary burrows constituted 5.6%; 47.3% 
were recently dug and had from 2 to 10 exits; 29% of the burrows 
were of medium age and had from 1 1 to 25 exits and 18.1% had from 
26 to 80 exits; these old burrows were, presumably, dug decades ago. 
More than 70 exits in one burrow are known in Kanin, up to 60 in 
Lena-Khatanga territory, up to 40 in the Commander islands and up 
to 30, in the north of the Taimyr peninsula. It happens that two or 
more families may occupy one burrow. On the Commander islands, 
arctic foxes inhabited one burrow for 15—20 years (Cherskii, 1919— 
1920; A. Romanov, 1941; Il'ina, 1949-1950; Sdobnikov, 1958; 
Skrobov, 1958; Maksimov, 1959). 

In level places, or among low hills, the area of intercommunicat- 
ing passages, representing a whole burrow system not rarely occupies 
a territory of 100-200, and sometimes 500 m^ On Belyi Island, 
burrows were found in the form of systems of communicated pas- 
sages with 53 exits occupying 1800 m^; during the year of this 
observation, 11 exits were made. Along the Tiutei river in Yamal, 
more than 10 burrows were close to each other. On Belyi Island, in 



322 




; 02 xj /of'^ %oo5 



Fig. 86. Scheme of the construction of an old arctic fox burrow (from A. 

Romanov, 1941). 
1 — primary holes, 2 — secondary holes, 3— nest, 4 — underground passages, 

5 — latrine. 

an area of 3500 m^, 8 burrows were found. On Wrangel' island, the 
colonies of arctic fox occupy entire sandy hills and are riddled with 
countless passages (Mineev, 1935; Tyulin, 1938; N.N. Spitsyn). 
227 In old burrow systems, arctic foxes make passages in 2—3 galler- 
ies with some chambers for nests. The den in which the nest is placed 
is lined with dry grasses and moss. The size of the nest chamber is 
from 0.5 to 1 m in diameter and 30-50 cm in height. The diameter 
of the exit opening of burrow ranges from 15—30 cm, rarely more, 
reaching in particular cases 80 cm. The cross section of the under- 
ground passages is 12—20 cm in height and 15—30 cm in width. As 
a rule, the horizontal cross section of the passages and the exits 
somewhat exceeds the vertical (Cherskii, 1919—1920; Boitsov, 1937; 
Tyulin, 1938; Dubrovskii, 1940; Chirkova, 1940; A. Romanov, 1941). 
The area occupied by the burrows, the underground passages and 
dens are very dirty because of the remains of food and excrement. As 
a result of plentiful manuring of the soil with feces and remains of 
food, rich dense dark green vegetation, up to 0.75 m high (in Kola 
peninsula) develops. Plant formations on the burrows are highly vari- 
able and represent a type of northern herbaceous vegetation. In such 
"arctic fox meadow gardens", tens of higher plant species grow which 



323 

are absent in the majority of tundra associations. Dandelion, polar 
poppy, Alpine forget-me-not, and foxtail grow in Taimyr below the 
latitude 75° N. Lat. Near the entrances to burrows mosses {Mnium, 
Drepanocladus) grow. Burrows that were occupied by arctic foxes in 
previous years are characterized by camomile [Matricaria], meadow 
bluegrass [Poa pratensis] and others, or by dense herbaceous growth 
mixed with fire-weed and bistort. On abandoned burrows, often only 
short fescue [Festuca supina] is found (Dubrovskii, 1937; Zolotov, 
1940; Sdobnikov, 1958; Maksimov, 1959). From a distance, it is easy 
to determine the location of burrows, which is evident in level tundra 
from 0.5 km, and with binoculars, up to 1.5 km. From aircraft, bur- 
rows can be seen well from a height of 400-500 m; and from 1000 
m, it is possible to map a majority of centers of arctic fox burrows 
(Zolotov, 1940; Freiberg, 1940; A.F. Chirkova). 

In winter, in periods of continuous snow storms, arctic fox makes 
burrows for shelter from driving snow on lee sides. The entrance is 
blocked by the blowing snow and the animal remains in the burrow 
until clear weather returns. Snow burrows are not longer than 1—1.5 
m, widening at the end, of dimensions [width] 25-40 cm and height, 
25 cm. The arctic fox also builds temporary burrows near food ac- 
cumulations thrown up by the sea (Nosilov, 1909; A. Romanov, 1941 
and others). 

In years of massive reproduction, when all littering burrows are 
occupied, arctic foxes carry the pups outside the burrows — some- 
where under the shelter of grasses, where they stay under cover of 
grasses, in a pile jof flotsam, or even in the hollow of a large trunk 
of the latter (Romanov, 1941). 

Daily activity and behavior. The arctic fox is active at any time 
of the twenty-four hour period. Scarcity and variability in the food 
supply force the animal to wander widely in search of food. In autumn 
and in winter, the arctic fox feeds mainly at night; in spring and in 
summer, apparently, in connection with the beginning of breeding and 
the appearance of their young — during the whole 24 hours. The arctic 
fox frequently rests in midday; in clear, windless weather, it settles 
down anywhere on a tussock, hillock or slope protected from the 
wind, on ice-hummocks or it buries itself in a snow-drift. In spring, 
and in the first period after the (re)appearance of the sun, the animal 
lies by a dark cliff or on a precipice where it is well warmed. In the 
Lena-Khatanga region, a female arctic fox rested in June after eating 
from 15:00 to 17:00 during the day and after that from 21:30 in the 



324 

evening; a male, in October arose from rest at about 06:00 in the 
morning and retired again at 14:30. On Mednyi Island, in good weather 
in the middle of June, a whole family of arctic foxes may be seen near 
the burrow early in the morning and again after 17:00 (Cherskii, 
1919-1920; A. Romanov, 1941). 

While sleeping, the arctic fox rolls itself into a ball, buries the 
nose in the base of the tail and covers the belly and limbs with the 
tail. Arctic foxes sleep deeply; during periods of noise — ^wind or ice- 
motion — ^it may be caught by hand (Mikhel', 1937; Rutilevskii, 1939). 
This is easily done during times of hunger, when the arctic fox is 
seriously weakened. 

The arctic fox is less sensitive to fluctuations in temperature than 
red and corsac foxes, as a result of which, at positive [above-freez- 
ing] temperature, the general activity of the arctic fox is less than at 
228 negative. By lowering of temperature from +15° to + 5°C, the fre- 
quency of breathing of the white fox is increased. In the blue fox, 
within the limits of 0—3 0°C, such dependence is not observed. Gas 
exchange and heat production are greater in summer and less in 
winter. Daily heat production for 1 kg body weight is 450 kilo- 
joules^', in autumn, it is higher and in spring and especially in 
summer, it is still higher, reaching 579 kilojoules (Kalabukhov and 
Poluzadova, 1946; Kalabukhov, 1950; Pozdnyakov, 1953). 

In spring and summer, arctic foxes are most active on the quiet, 
cloudy days and in warm weather, with light wind. On cold windy 
days, and especially during heavy rain, its activity is shaфly reduced, 
and the animal conceals itself in a comfortable place. In winter, arctic 
foxes do not go out during strong snowstorms, in freezing weather 
with severe wind, and on dark nights. On the contrary, they are highly 
active on clear moonlit nights, during northern lights and at dawn. 
Arctic foxes do not react to light wind, but in a strong one they move 
into it. Upon encountering wind, they come to a halt, most commonly 
in mild weather. On Belyi Island, they fall in traps usually in calm 
weather and during weak and moderate wind speed of 6—7 m/second. 
On the Chukotka peninsula, arctic foxes, on the contrary, are more 
active and incautious during violent snowstorms and frequent inspec- 
tion of the traps at this time yields a large catch (Freiberg, 1934; 
Dubrovskii, 1937; Tyulin, 1938; Shundik, 1948; Yanygin, 1948; 
Sdobnikov, 1958). On the Solovetsk islands, up to 50% of the arctic 

-^'According to international system of units (IS). 



325 

foxes go seeking for food at a right angle to the wind, up to 40% go 
with the wind and only 10% go against the wind (Boitsov, 1927). The 
same is observed on the Murmansk coast. 

As compared to the red foxes, the arctic fox is bold and more 
trusting. Steller (1774) mentions how "innumerable packs" of arctic 
foxes surrounded a camp of people and became each day "more 
courageous and mischievous". The arctic foxes strewed about not 
only provisions which were even stored in barrels but also invaded 
people's living sites, chewed and dragged off clothing, shoes, and 
instruments. During his journey south from latitude 85° N. Lat., F. 
Nansen (1897) was also subjected to "attack" by a great number of 
arctic foxes which strewed about his belongings. Now, arctic foxes 
are significantly more cautious. Nevertheless, factory workers in 
Yamal, Taimyr and others have more than once reported the pillage 
of the meat and fish stores and other products by arctic foxes. When 
an arctic fox is driven with reindeer, the exhausted anim.al hides 
under a fur thrown on the snow. Arctic foxes on Ushishir Island are 
very unwary (Kuril chain; Klumov, 1960). 

Mousing animals and also migrating arctic foxes frequently no- 
tice humans only at a distance of several meters. In this case, the 
suфrised animal sits down, looks at the person, sometimes stretches 
and then jumps up and continues on its way. On the Commander 
islands, where arctic foxes are protected, they inhabit villages, 
bringing their pups near houses or sheds. The parents, usually in turn, 
keep close to their litter, and after becoming accustomed to people, 
take the food from their hands. They may wander for hours only a 
few steps behind people, barking and sometimes sniffing inhabitants 
(Cherskii, 1919-1920). 

In captivity, arctic foxes quickly become tame. Getting accus- 
tomed to humans, some arctic foxes return after being set free. In the 
arctic fox, conditioned reflexes are easily developed. On the Com- 
mander islands, the animals knew well, by the clock, the time of their 
supplemental feeding and quickly respond to a sound signal. To a 
whistle, accompanied by the distribution of dried salmon or fur-seal 
meat, "arctic foxes appeared as if from underground, ran immedi- 
ately, and yelping jumped on the handout" (Barabash-Nikiforov, 1937). 

The relationships of the arctic fox to various animals are diverse. 

Above were mentioned reindeer and polar bear. Flocks of brant geese, 

gulls and skuas actively repulse the attack of the predator. On the 

229 Commander islands, the arctic fox drives pigs away from the flotsam. 



326 

but at the same time, it plays with tethered goats. Arctic foxes are 
afraid of cats and dogs, especially where people incite them. In Novaya 
Zemlya. arctic foxes follow dogs to carrion, and sometimes, espe- 
cially the young, play with them and even with humans (Shrenk, 
1855; Nosilov, 1909; Cherskii. 1919-1920). 

On a level surface, a dog can easily catch the arctic fox. but in 
hilly or tussocky places or in ice hummocks, the arctic fox runs away 
from it. The manner of the arctic fox is similar to that of the corsac 
fox; it runs in zigzag with sudden sharp turns; it confuses dogs. 

In catching lemmings, the arctic fox waits for a long time for the 
appearance of a rodent from a burrow, then jumps and crushes it. 
Listening to the squeak or rustle of a lemming, the arctic fox can in 
a few seconds excavate the snow to a depth of 1/2 m or more. \\Ъ11е 
hunting for lemmings, the arctic fox can easily come within gunshot 
of a man. A case is knov^n of an arctic fox which, while digging up 
small rodents during a snow storm, was caught by a hunter by the tail. 
On seeing a gull on a rock on the shore from a distance of 120 
meters, an arctic fox not yet molted went on with slow steps, confin- 
ing itself to patches of snow; at 15—18 m, it lay and crept on its belly, 
and then with a strong jump caught the gull (Chastin, 1939). At the 
molting time of geese, arctic foxes hunt them communally, driving 
the helpless bird to their comrades; the arctic fox throws the captured 
goose on its back and pulls it. It drinks the eggs of large birds through 
holes made by the canines. 

On the Commander islands, wandering constantly among the 
hauling places of fur seals, arctic foxes search for not only dead 
animals, but also pull away the yoimg fur seals from lone females 
holding them by a unique method, by the lips; they take mainly the 
sick and weak (Barabash-Nikiforov, 1937). 

The arctic fox eats part of a large prey on the spot, and carries 
the remainder to a burrow or buries it. In years with abundant food, 
the arctic fox makes a store, collecting some lemmings, fish or large 
birds; it buries their bodies in the ground or in sand, firmly tamping 
the surface with its nose; it pushes them imder stones, ice blocks and 
so on. A case is known on Mednyi Island when arctic foxes took 
about hundred eggs from the tin box of some sleeping hunters, 
bur>ing the greater portion in a laida. On the same island, they 
obser\'ed the return of arctic foxes to their stores to eat them (Cherskii, 
1919-1920). AU the same, a great portion of the hidden food is not 
utilized bv arctic foxes. 



327 

The behavior of the arctic fox changes in various years depend- 
ing on the abundance of food, prevalent weather and other reasons. 
With abundance of lemmings, white hares and willow ptarmigan, the 
arctic fox in the Bol'shezemel'sk tundra wanders little, and restricts 
itself the whole winter in proximity to its burrows, frequently hiding 
in them. When feeding on a second grade food [source], arctic foxes 
behave uncomfortably, but nonetheless do not leave summer habitats. 
In case of food deficiency, the animals go to the seashore or migrate 
in other directions (Skrobov and Kozhevin, 1957). In years of famine, 
arctic foxes move close to the winter stations and villages, lose their 
prudence and even defend their prey against humans (Commander 
islands, Tomsk district) (Cherskii, 1919-1920; Likhachev, 1930). 

The arctic fox takes to water and during migration they swim 
across large lakes and rivers, for example, the lower Taimyr, 400 m 
in width and a current velocity of 3.5 m/s. They swim across the 
lower Khatanga (1-1.5 km) as well as Yenisei and Ob', straits in the 
Bol'shezemel'sk tundra (2 km wide) and to Kil'din Island (Tret'yakov, 
1871; Formozov, 1929; Rutilevskii, 1939; Shastin, 1939; Chirkova, 
1955 and others). In Alaska, the arctic fox has mastered straits up to 
4 km wide. 

The speed of the arctic fox in water is 2-2.5 km/h, slower than 
in the dog. While swimming, the animal holds its back and tail above 
water, the latter at a small angle. When the tail gets wet, indicating 
fatigue, the fox quickly drowns (Shastin, 1939). The arctic fox swims 
also across rivers containing slush at water temperature of about 0°C; 
on being turned around several times in a whirlpool, the arctic fox 
230 does not become disoriented and swims in a defined direction. On 
coming out on shore, it shakes itself and immediately runs rapidly 
away from the shore. 

At the beginning of migration, masses of arctic foxes gather by 
rivers and run at the very edge of the water (see also p. 330). As long 
as ice is smooth, arctic foxes gather on the lakes, playing and harry- 
ing one another. At the sound or sight of an а1ф1апе flying at a height 
of 100-200 m, the arctic fox takes a defensive pose and lies on its 
back (V.P. Makridin); at such a time, the red fox runs away. 

On the Commander and Solovetskii islands, according to the 
behavior and hunting habits, arctic foxes are divisible into the follow- 
ing groups: 1) Sedentary animals, leading an inactive way of life, and 
living in permanent places near human settlements, chiefly by scav- 
enging; 2) half-sedentary animals, carrying on in a region of permanent 



328 

habitation, but sometimes moving to places inhabited by neighboring 
groups; 3) wandering arctic foxes. On Mednyi Island, the more alert 
and active are the tundra arctic foxes who hold aloof from people. On 
the Solovetskii Islands, vagrancy is developed mainly among males. 
The predominance of males among the migrating animals is, appar- 
ently, characteristic of arctic foxes of the mainland tundras 
(Barabash-Nikiforov, 1937a; Boitsov, 1937; Chirkova et al., 1959). 

The behavior of the arctic fox varies shaфly with the beginning 
of the reproductive period (Cherskii, 1919-1920; Barabash-Nikiforov, 
1937a; Boitsov, 1937; Chastin, 1939; A. Romanov, 1941 and others). 
On Mednyi Island, the premating roll-call among males starts early 
in January. At night in February, the plaintive howling, yelping, and 
squealing of females are heard. In March males frequently urinate 
near burrows and feeding places; females urinate, raising the tail, and 
allowing males to approach them. The animals run, often holding 
something in their teeth — a bone, stick or lump of snow. Playing 
between mated pairs alternates with hard fights among males. In the 
location of the fight, bits of wool, and sometimes blood are left; on 
Mednyi Island, it was observed that at that time, arctic foxes injure 
the genital organs of each other. In fights, up to ten animals are 
involved. In the northern Taimyr peninsula, nuptial games were ob- 
served on 20 April 1936. The mating of arctic foxes is accompanied 
by loud screeches. 

In some instances, male and female become strongly attached to 
each other in the breeding period. On Mednyi Island, a female which 
had been taken from her chosen male and kept in an open-air cage 
with another male, lived with him the whole spring, but did not mate 
with him. During the same period, she always expressed her attention 
to the first male, who sometimes passed by. The latter, although he 
had his own mate, brought food to the caged female. 

During pregnancy the males constantly offer food to their fe- 
males and with special diligence not long before the appearance of 
the newborn. On Mednyi Island, a male carried fish ''my agon 'ku" 
{Cyclopterichtys venticosus) to a female that happened to fall captive. 

Shortly before parturition, the behavior of a captured female 
becomes more quiet and cautious. With the appearance of the babies, 
both parents feed the young in turn, bringing them food and playing 
with them, but most often the mother remains near the burrow to 
guard the litter. In case of danger, the female sometimes hides in the 
burrows together with the young while the male strives to drive away 



329 

the person, howling and gradually moving away from the litter. In the 
absence of adults, the young allow a human to approach within sev- 
eral paces, and even after fleeing to the burrow, they appear again 
after a few minutes. At the age of 3-4 weeks, the animals leave the 
burrow, on Mednyi Island already at the end of June and the middle- 
end of July, they start playing independently. Making excursions, 
they frequently change burrows. At this time, their barking is first 
heard. In Lena-Khatanga territory, the young arctic foxes start to play 
within 5 days after leaving the burrow, and within 10 days, they 
attempt to catch lemmings and excavate their burrows and begin to 
feed on meat. At the age of 1.5 months, they shift over to meat and 
23 1 other food. In the second half of July, they share with their parents 
in hunting molting geese. The mother feeds the young on milk for one 
to 2.5 months. 2.5—3 months after birth of the young, the father 
abandons the litter. At the end of their life together, the mother also 
leaves the litter for a whole day. After the female finally abandons 
the young, the litter breaks up. A case is known when, after the death 
of the father of one litter, the mother transferred the young to the 
burrow of another arctic fox, where both mothers fed the young 
together (Il'ina, 1949-1950). 

The more the year is deficient in food, the longer the time taken 
by the parents in search for food, and the earlier they leave the young. 
With food abundance, members of the family stay together until 
autumn, and separate gradually; individual arctic foxes live near 
burrows throughout the entire winter. In the former Kolymsk region, 
a female was found in a burrow together with her adult offspring, 
even in winter (lokhel'son, 1898). 

Seasonal migrations and transgressions. The arctic fox is a typi- 
cal migrant, and wandering is of great significance in its life. The 
migration of arctic foxes must be understood as a historically estab- 
lished phenomenon. If judged by Pleistocene-Holocene remains, this 
phenomenon has existed for many thousands of years (Tugarinov, 
1927; Gromov, 1928; Kirillov, 1950; Pidoplichko, 1951; Vereshchagin, 
1953, 1959, 1960; Serzhanin, 1961). 

The reason and stimulus for commencement of migration is in- 
sufficient food. This is confirmed by the appearance of massive and 
long-distance shifts of arctic foxes in years of the disappearance of 
the main food — lemmings. At the same time, the relationship between 
migration and food abundance is not invariant. The start of arctic fox 
movement away from the denning region is sometimes observed in 



330 

the presence of food. Migration is subject to instinct. In old animals, 
migration is also modified by experience. One may assume that the 
old arctic foxes react to signals announcing the coming of famine — 
an early cold snap with rain and wet snow, early departure of birds, 
the appearance of onshore and changeable winds and, possibly, other 
unknown factors. It was repeatedly observed that periodic migrations 
of arctic foxes from places of permanent habitation are repeated after 
2-4 years. 

Migrations of arctic foxes start in August-September, depending 
upon the time when food disappears in the main breeding places. The 
massive movement of animals when the arctic foxes "come in crowds", 
as it is said in the North, occurs in October— November. 

The route of arctic foxes before the freezing period runs along 
the water's edge, where a trampled path is formed. At Serebryanyi 
[Silver] bay on the western coast of the northern island of Novaya 
Zemlya, the whole shore was spotted with tracks of arctic foxes in 
1887 and the beaten tracks of animals from all sides met here. On the 
sands of the Khatanga river, in 1931, there was a path several meters 
wide (Nosilov, 1909; A.F. Chirkova). Part of the arctic foxes move 
on a wider front. In the northwestern Taimyr peninsula, arctic foxes 
move along in a 2—3 km deep strip along the shore. In the northeast- 
ern Bol'shezemel'sk tundra, the animals move along a front of tens 
of kilometers, but the majority of them are concentrated in the coastal 
zone of Baidaratsk bay (Kiфichnikov, 1939; Chirkova, 1955). 

In separate places, either latitudinal — ^western or eastern — direc- 
tions predominate, as in northern Arkhangel'sk district, or meridional 
ones, as the well-known route of the coastal arctic fox along the 
Yenisei river. Migrations with departure from the tundra to the forest 
zone are more noticeable, and early began to attract to themselves the 
attention of investigators. Meridional migrations are noticeable only 
when they are massive phenomena. In the neighborhood of Taimyr 
lake, the autumn migration in 1947 was in a southeastward direction; 
the arctic foxes swam across the lower Taimyr from the western 
shore (Sdobnikov, 1958). 

Animals penetrate the taiga zone in great numbers, going along 

the valleys of large rivers or coming out onto open watersheds; from 

the sea coast, they move out onto the ice and to the islands of the 

233 Arctic Ocean. The arrival of arctic foxes into the forest zone is 

frequently preceded by the flight of snowy owls [Nyctea scandiaca] 



331 

which more quickly respond to deficiency of food in the tundra and 
vacate it earlier. 

It has been observed that arctic foxes move as though in waves. 
Such migrations of arctic fox groups were observed in Lena-Khatanga 
territory. In the interior of the Bol'shezemel'sk tundra, in 1933/34, a 
movement from east to west began from beyond the Polar Urals, 
which was noticed at the end of October when in the course of a 
month there was a migration of well-fed arctic foxes. After a pause 
at the end of December, a smaller group of animals moved, and from 
January 20th, during the rest of the month, individual animals were 
seen running by on three days. The second and third groups of ani- 
mals were emaciated. On the sea coast, arctic foxes at the end of 
October moved the whole winter in the same western direction, the 
first strong wave occurring at the end of November, the next — ^at the 
end of December, and finally — in the middle of February and from 
the end of this month until the middle of March and later. The rut, 
which took place at the end of March and in April, did not stop the 
movement of arctic foxes, which continued running in the same di- 
rection as in winter (A. Romanov, 1941; N. Dement'ev, 1955). 

In winter, arctic foxes spread out over a huge territory of tundra, 
forest and ice landscapes. The total area inhabited by the species 
increases at this time by several times and at the same time popula- 
tion density in the tundra decreases. The majority of arctic foxes die 
in unaccustomed life situations, from hunger or diseases, and are 
caught by hunters. Long-distance migration of arctic foxes in the 
majority of cases is "a way to death". 

Those animals remaining alive in February— March, sometimes 
even in January, migrate back; in some years this extends until April 
and May. The first tracks of arctic foxes from the south appeared 
near Taimyr lake on the 20th of March, and the movement from the 
seashore into the depths of the tundra in 1948-1949 ended by the 
middle of May (Sdobnikov, 1958). Apparently, only a negligible 
portion of the arctic foxes return to the tundra. Individual arctic foxes 
remain during the warm period in uncharacteristic types of land- 
scapes like forests and on the ocean ice. 

Long-distance migrations are more often carried out by males (on 
the Solovetsk islands, up to 80%), but cases of long movements are 
also known in females, for instance in 1928, to the neighborhood of 
Krasnoyarsk. Long-distance movements seriously deplete arctic foxes 



333 

and suppresses their reproductive capacity (Sluchevskii, 1886; Skalon, 
1928; V.K., 1929; Boitsov, 1937; Chirkova, 1955, 1959 and others). 
In the forest zone, arctic foxes were observed and frequently 
hunted at 1000-1800 km, and in exceptional cases, 2000 km (for 
example near Komsomol' -na- Amur), and in the Arctic Ocean, at 
1000 km from the limits of summer habitation. The average daily 
route encompassed by the animal is, apparently, about 20-30 km. 
234 At the end of the 1930's, 172 arctic foxes were marked in the 
Novyi Port region on Yamal, of which 3 1 animals were re-captured 
(Sdobnikov, 1940; Fig. 88). A maximum translocation of about 1200 
km in a straight line as it went across ocean ice to the north island 
of Novaya Zemlya, was made by one animal. The majority of arctic 
foxes moved off in a southwesterly direction; the farthest ones were 
caught near the village of Muzhi on the Ob', at the mouth of the 
Pechora, at the Snopa river on Chesh Bay, at the southern end of 
Novaya Zemlya, and at its northern limit. 

When food decreases in years of the animal's abundance, almost 
all arctic foxes migrate or disperse to different areas away from their 
denning places. Districts to which they dispersed to the south for the 
majority of migrating arctic foxes were to be found in the northern 
subzone of the forest. When a peak of migration is reached along a 
permanent route, one may observe large numbers of foxes. Thus, in 
1887 during one stroll on the western shore of Novaya Zemlya, 
Nosilov (1909) encountered hundreds of animals, and in the course 
of two weeks in October, by his estimate thousands of foxes passed 
by. After migration to the southern island, only single individuals 
remained. On the southern coast of Belyi Island at the end of Septem- 
ber 1930, tens of foxes were seen, through binoculars, wandering 
about awaiting the formation of fast ice (A.F. Chirkova). On Dikson 
Island, on good "movement" days 5-6 arctic foxes were seen directly 
in a distance of 1 km (Heptner, 1936). On the lower Taimyr river, 
between 18 and 20 September 1935, each day up to 21 foxes swam 
across while on September 1947, 10 to 15 foxes were observed at one 
time within a radius of 2 km in this region (Sdobnikov, 1958). On the 
Chelyusk peninsula, during the time of the arctic foxes passing at the 
end of October, one might see at one time up to 46 animals, after 
which the number dropped shaфly; during winter, there were only 
separate individuals. The total number of foxes that passed through 
an area of 10 km in autumn 1932 constituted not less than 1500 to 
2000 individuals, and in spring, during the period of their return, 



334 




233 Fig. 88. Directions and limits of migrations of individual arctic foxes marked on 
Yamal (from Sdobnikov, 1940): 1 — direction of movements; 2 — no movements. 

there were not less than 800 (Rutilevskii, 1939). On the right bank of 
the lower IChatanga, upstream from Cape Bol'shaya Korga, tens of 
passing arctic foxes were counted each day on 24 and 25 August 
193 1. Along the left bank of this river near the mouth of the Novaya 
river, from 2 to 80 fresh tracks of arctic foxes were noted on a 10- 
km route at the beginning of October, and below Khatangsk village, 
it was possible to observe 2 to 4 migrating arctic foxes at one time 
at any time of day starting from September 17; after ice formation on 
October 10, arctic foxes disappeared. In that period, no less than 1000 
animals passed by Khatangsk village (A.F. Chirkova). 

In Arkhangel'sk district, judging by the number of prepared skins, 
the number of arctic foxes emigrating for long distances only in a 
south and southwestern direction constituted, in some years, about 10, 
20, or 30%. No less than this number of arctic foxes departed to 
the north across the ocean ice. In this manner, during years of exten- 
sive migration up to 2/3 of all arctic foxes abandon the tundra zone. 

At the time of massive movements, the behavior of arctic foxes 
changes. At the beginning of migration, the direction of movement of 



335 

individual animals is variable, undefined and might be directly oppo- 
site one another. In this time, arctic foxes wander nervously and swim 
across rivers and canals. Coming to the banks of large and as yet 
unfrozen rivers, groups of arctic foxes start to change direction along 
them, running along the sea coasts and along mountain ranges and 
valleys; they move in response to the density and depth of snow 
cover, the weather, especially winds, and also to the availability of 
food along the way. 

At the beginning of migration, arctic foxes move in pre-twilight 
or early morning accustoming themselves to the trails trodden by 
previous confreres; without looking aside they hurriedly overcome 
the difficulties along the way. Later arctic foxes energetically run, 
trot or even gallop day and night, with this frequently barking or 
howling. Being distracted for a short time to search for food, their 
hunger barely satisfied, the animals again start on their predetermined 
way. 

On clear moonlit nights or during the period of the northern lights 
[aurora borealis], active movement is initiated. On their route, the 
arctic foxes move through villages, especially at night, and it is only 
during the light part of the 24-hour period, when people and dogs are 
present on the shore that the animals detour or lie in wait for the 
235 departure of people. Their instinct for wandering is so strong that the 
animals may stop for some time only to spread out at the beginning 
of migration to feed up, although the animals still remain in the 
denning region or have only started to migrate. During the peak of 
their movement, hunting for food detains the animals only briefly. 

T^rctic foxes from different geographic populations may sometimes 
mix during migration. Thus, in the winter of 1956/57, in a year of migra- 
tion from the east, an average of 50% of the arctic foxes captured in the 
vicinity of Клга at the very edge of the Bol'shezemel'sk and Yamal 
tundras were relajted by their fiir to the Obdorsk type, characteristic of the 
Yamal foxes. In the westem Bol'shezemel'sk tundra, such animals con- 
stituted 30%, and on the Kanin peninsula, less than \0%; all those 
remaining were referable to the local Pechora type characteristic of the 
European north (Chirkova et al., 1959; Fig. 82). 

Reddening of the fur" that is observed in arctic foxes and 
other carnivores on the underside of the paws, rarely in the axillary 

"In microscopic view, the reddish color stains equally every hair. The color is 
not soluble either in water or in soap, and it resists the action of alcohol, ether, 
chloroform, xylol and hydrogen peroxide (Chirkova et al., 1959; Shilyaeva, 1961; A.I. 
Solntsev). 



336 

depressions and on the belly, is obviously caused by skin gland se- 
cretions connected with periods of great stress on the organism, and 
appears particularly intensely during the time of the animal's migra- 
tion. Thus during the extensive migration of arctic foxes in the 
northeastern Bol'shezemel'sk tundra in the autumn and winter of 
1951/1956, the reddening of the fur of the foot pads and between the 
toes was noted in approximately half of the arctic foxes captured in 
the Kara region (42% bright color). In the following year during the 
absence of migration, reddening of the paw fur was only noted in a 
small number of animals (2% bright color in all) (Chirkova et al., 
1959). 

Basic migration routes and long-distance movements of arctic 
foxes are known over a large part of the species' range (Fig. 87). On 
the Kola peninsula, the arctic fox migrates mainly along the valleys 
of Kamenka, Kochkovka, and other rivers, [flowing] from the interior 
of the peninsula to the sea. From the northwestern Kola peninsula it 
runs southward to Murmansk and farther, along the railway line. 

Rare cases of the appearance of individual foxes were observed 
in the territory of the Lapland preserve and on the Tersk shore in the 
Olenitsa and Kuzomen' regions (Ozeretskovskii, 1904; Dubrovskii, 
1939; Zolotov, 1904; Nasimovich et al., 1948; Drebentsov, 1959). 

After the formation of fast shore ice and the appearance of ice 
at the mouth [entrance]* of the White Sea, the arctic fox moves off 
to the Kanin side, from whence they return in March, before the start 
of ice breakup. The large exodus of the arctic fox is repeated every 
4 to 6 years. The emigration of arctic foxes to the Varanger Fiord 
side, and the event of their appearance in northwestern Finland near 
Enonteklossa is notable (Юеето1а [and] Laine, 1961). 

Counter-movements across a wide distance take place in the 
European north, to the east of the White Sea, from the northeast to 
the southwest and west through the Bol'shezemel'sk, Malozemel'sk 
and Timansk tundras (Folitarek and Chirkova, 1930; Sergeev, 1934; 
A. Korsakov, 1941; G. Naumov, 1941; Maksimov, 1945; Chirkova, 
1955). Arctic foxes skirt to the south of Chesh, cut across the Kanin 
peninsula or follow around it, turning to the north along the eastern 
shore, and drop down to the south along the western; they migrate 
along the Zimnii coast of the White Sea to Dvina Bay. The arctic fox 



"The Russian word used is gorla, meaning throat — Sci. Ed. 



337 

frequently crosses the lower Pechora and other large rivers in specific 
places. 

Many arctic foxes migrate southwards along the Pechora, Mezena, 
Pinega, Northern Dvina, Onega rivers and their tributaries to the 
upper Vychegda, Sukhona, Yuga and others. The most extensive 
emigrations of the arctic fox, up to 58—56° N. Lat., were known even 
in the past century, and also in the present century in southeastern 
Latvian SSR, in Pskov, Novgorod, Leningrad, Kalinin, Vologda and 
236 Kostroma districts, and in the northwest and east of the Kirov district. 
Migrations were noted as far as the vicinity of Troitsko-Pechorsk 
village and the mouth of the Ilych river, and also in Ust' Kulomsk, 
former Storozhevsk and Letskii regions (60° N. Lat.) of the Komi 
ASSR (Middendorf, 1869; Sluchevskii, 1886; Dmitrieva-Sulima, 1911; 
Paramonov, 1929; Ognev, 1931; Lavrov, 1932, Sergeev, 1934; Guber, 
1937; Ostroumov, 1949; S.V. Shibanov, A.F. Chirkova). 

In the Pechoro-Ilych preserve, arctic foxes during the period 
from 1929 to 1949 appeared in the course of 9 winters: in 1929/30, 
1935/36 and 1937/38 there were massive migrations, and in 1933/34, 
1938/39, and in the course of four winters from 1944/45 to 1947/48, 
individual animals were encountered. Apparently, movement of the 
arctic fox on the territory of the preserve was limited by the depth of 
snow cover; in years of large transgressions, the average maximum 
depth of snow cover was 74 cm, but, in winters of occasional appear- 
ances — ^98 cm. In heavy snow years, the arctic fox often remained 
along rivers, moving on the ice, being malnourished and diseased 
(1929/30 and 1935/36). In light snow winters (1937/38) they moved 
freely through the taiga. There is a supposition that occasional arctic 
foxes, moving into the preserve in 1944—1945 continued to live there 
and did not depart for 4 years; after the capture of two animals in 
1947/48, no more foxes were encountered (Teplov, 1960). 

Some of the arctic foxes from the northeastern Bol'shezemel'sk 
tundra move over to Vaigach and Novaya Zemlya, across the ice of 
the Barents Sea, and then arrive at Kolguev [island]. The migration 
of arctic foxes in Novaya Zemlya was intensive in the last century 
along the western and eastern shores of both islands (Nosilov^ 1909). 
Good passages of arctic foxes on Novaya Zemlya occur twice a 
year — ^in autumn and spring (Dubrovskii, 1941). In different years, 
arctic foxes move from Bol'shezemel'sk tundra to the Urals. Waves 
of massive migrations in the north of Arkhangel'sk district were 
noted in the winters of 1909/10, 1914, 1926/27, 1929/30, 1935/36, 



338 

1937/38, 1944/45, 1946/47, 1947/48, 1954/55, 1956/57 (Dmitrieva- 
Sulima, 1911; Folitarek and Chirkova, 1930; Bashkin, 1940; 
A. Korsakov, 1941; G. Naumov, 1941; Maksimov, 1945; Chirkova, 
1955; Chirkova et al., 1959). 

Basic migration routes on the Yamal peninsula, when, accord- 
ing to Nentsi stories, foxes run "in herds," go in a meridional 
direction, along the shores of Ob' Bay and the Kara Sea, and also 
along the central Khoi watershed. The arctic foxes move out over 
a wide front along the ice of the Kara Sea. 

Less significant are migrations along the Taz and Pur rivers to 
the east on the Yenisei. The most southern places in west Siberia 
where arctic foxes appear are noted in the vicinity of Nyaksimvol' 
(63° N. Lat.); in the belt of southern taiga along the course of the 
Irtysh and Tobol, below Tobolsk and even Yalutorovsk (56.5° N. 
Lat.), below Tevriz (57.5° N. Lat.). Dispersal of foxes to the vi- 
cinity of Tom'sk (57° N. Lat.) is known (Skalon, 1928; Formozov, 
1935; Andreev, 1937; Laptev, 1958; M.A. Sergeev). Long-distance 
movements of arctic foxes into the forest zone of west Siberia 
took place in 1921, and in the seasons of 1926/27, 1929/30, 1932/ 
33, 1937/38, 1941/42, 1944/45, 1947/48, 1954/55, and less 
significantly in 1930/31, 1933/34, 1935/36, 1939/40, 1946/47 
and later. Migrations of the arctic fox are also known in 1910 and 
1913. 

In northern Krasnoyarsk territory, the arctic fox takes itself 
upstream along the Yenisei to the mouths of Lower and even the 
Podkamennaya [stony] Tungusk. Occasional arctic foxes reach 
Krasnoyarsk (V.K., 1929, and others), the upper course of Lower 
and Podkamennaya Tungusk (60° N. Lat.), to the region of the city 
of Bratsk, even the villages of Aban and Taishet along the main 
Trans-Siberian railway (56-57° N. Lat.). From the Taimyr peninsula, 
large numbers of arctic foxes migrate along the Pyasina and Nizhnaya 
Taimyr rivers and also along the shore of Taimyr lake, departing from 
the tundra zone through the belt of northern taiga forest to the Central 
Siberian Plateau. A well-defined migration of arctic foxes takes 
place along both shores of the Khatanga gulf and Khatanga river 
and its tributaries, the Popigai, Kheta and Kotui, where the arctic 
fox reaches the upper courses of these rivers. To the north, arctic 
foxes disperse across the ice to all the nearby islands, in particular 
Sevemaya Zemlya and Vize island — ^at 78° N. Lat. 



339 

Large migrations and long-distance dispersal of arctic foxes in 
the northern Krasnoyarsk territory are known in 1860, 1913, 1922/ 
237 23, 1925/26, 1927/28, 1931/32, 1935/36, 1944/45, 1946/47, 1947/48, 
and 1954/55 (Tret'akov, 1871; Aleksandrov, 1927; Buryakov, 1929; 
N. Naumov, 1930; Heptner, 1936; Shastin, 1936, 1939; L. Popov, 
1939;Chirkova, 1955). 

Arctic foxes in Yakutiya wander not only along the rivers but also 
in the interfluve country along the Kharaulakh, Chekanovsk and other 
ranges. Transgressions were observed in the upper Anabar, Olenek, 
Vilyui, and Lena rivers (Mukhtuya and Olekminsk villages, 61° N. Lat.) 
and along the Vitim (Mama village, Irkutsk district, 58° N. Lat.), in the 
vicinity of Yakutsk and Krest-Khal'dzhai village on the Aldan. Arctic 
foxes were found along the Omoloi and Yana rivers to the latitude of 
Verkhoyansk, along Indigirka to Oimyakon village (63° N. Lat.), on the 
upper Kolyma to Seimchan (62.5° N. Lat.) and its tributaries, the 
Yasachnaya and Korkodon (Tkachenko, 1932; Tugarinov et al., 1934; 
Kolyushev, 1936; Guber, 1939; Skalon, 1940; Skalon et al., 1941; A. 
Romanov, 1941; Belyk, 1953). Extensive migrations of foxes take place 
here in the tundra zone and along the shores of the Laptev and East 
Siberian seas, in both easterly and westerly directions. Arctic foxes 
come to the Anabar river basin following the migration of wild rein- 
deer herds from the eastern Taimyr. 

Commonly, arctic foxes disperse across the ice of the Arctic 
Sea, to the New Siberian and other islands (V.U., 1935, Zarovnyaev, 
1933; A. Romanov, 1941, and others). Masses of arctic foxes, 
together with seals and polar bears, remain in winter among the 
[ice] hummocks in the sea 250 to 350 km from the mouth of the 
Olenek river; here these animals seek their living. After strong 
winds and snowstorms, the arctic fox wanders back to the sea- 
coast (Vasipev, 1935). 

Migrations of arctic foxes in the western and eastern regions 
of the Yakutsk ASSR do not proceed simultaneously. Large migra- 
tions of arctic foxes to the east of the Lena (along the Kolyma, 
Indigirka, Alazeya and others) were observed in 1895, 1903 and 
1944/45; and in the western regions, in 1926/27, 1932/33, 1947/48, 
and 1954/55. In 1929/30 and 1935/36, long migration of arctic foxes 
originated, apparently, on the whole territory of Yakutiya (lokhel'son, 
1898; Buturiin, 1913, E. Sh., 1930; Tugarinov et al., 1934; Skalon, 
1940; A. Romanov, 1941; Chirkova, 1955). 



340 

F. Vrangel' (1841) even saw arctic foxes on the ice of the East 
Siberian Sea between the Medvezhie islands and the De Long 
strait. Some information is available on movements of arctic foxes 
along the shores of Chaun bay to Wrangel' island and back at Cape 
Uelen, and eastward across the Bering Strait, and also in a south- 
western direction along the shore of the Bering Sea (Mineev, 1935; 
Chirkova, 1955 and other materials). 

Transgressions of arctic foxes are rarely observed on 
Kamchatka — in the Parapol valley region on the isthmus, on the 
western coast to Ust'-Bol'sheretsk, the southern end of the penin- 
sula and the northern Kuril Islands; on the eastern coast of the 
peninsula — ^to Ust'-Kamchatsk and the Kronotsk peninsula. In 1895 
and 1896, several arctic foxes were caught near Petropavlovsk 
(Paramonov, 1929; Sergeev, 1936; Guber, 1939; Yu.V. Averin). 

Arctic foxes were noted at Gizhiga and Penzhino (Ognev, 1 926) 
and along the western shore of the Okhotsk Sea (Sergeev, 1936; 
Guber, 1939). Nordmann (1861) was the first to point out the oc- 
currence of the arctic fox in the lower Amur. Later transgressions, 
or possibly the transporting of separate individuals on ice, were 
observed at 51-52° N. Lat. on the western shore of the Tatar strait, 
in the Lll'chsk region in the winter of 1949/50 and in the vicinity of 
Komsomol' sk-on- Amur in the autumn of 1958 (Belyaev, 1959). 

F. Nansen encountered arctic foxes on the ocean ice below 85° 
20' N. Lat. in April 1895, 300 miles from the nearest land (Franz Josef 
Land). Brusilov (1934) during the period of the "Northern Агшу" drift 
[ship in ice] noted the presence of arctic foxes in the winter of 1913/ 
14 on the ice between the islands of Novaya Zemlya, Franz Josef Land 
and Sevemaya Zemlya, approximately from 77° to 82° N. Lat. and from 
60° to 80° E. Long. De Long (1936) saw arctic foxes and their tracks 
at a latitude of 72° to 77° N. Lat. between Wrangel', Geral'd and Bermet 
islands. During the voyage of the ice-breaker "G. Sedov" and later on 
238 the drifting polar stations "Sevemyi Polyus" [North Pole] 2, 3 and 4 
in the "inaccessible pole" and other parts of the ocean, arctic foxes 
were found between 77°-87° 45' N. Lat. and 129° E. Long.-157° W. 
Long, at distances ranging from some hundreds to 1000 km to the north 
and northeast of the New Siberian islands (Badigin, 1940; Buinitskii, 
1945, 1946; Karelin, 1945; Ostrekhin, 1945; Chapskii, 1946; Rutilevskii 
and Uspenskii, 1957; Kirillov, 1958). The most northem point at which 
the arctic fox has been observed is -88° N. Lat., at a distance of 800 
km from the nearest land — ^Ellesmere Land (Uspenskii, 1956). The 



341 

majority of occurrences of arctic foxes met with on ice are noted in 
November-December, and rarely in March-April. 

In the Westem Hemisphere, arctic foxes migrate also in various 
directions, both to the south and to the ice of the Arctic Ocean. On the 
eastem Atlantic coast of the North American continent, arctic foxes 
descend along the treeless regions significantly farther south than on 
the westem Pacific [coast]. Up to 90% of the arctic foxes within the 
arctic archipelago spend the winter on ice. Arctic foxes even disperse 
from Canada to Greenland. 

Reproduction^^. Animals begin to reproduce at the end of the 
first year of their life, although they attain full maturity in the 
second year. Development of the reproductive organs begins with 
increase of the testes in autumn. In a year with unfavorable con- 
ditions, the formation of spermatozoa and their appearance in the 
excretory ducts in adult wild arctic foxes of Yakutiya, began in 
February at a testes weight of 2600 mg and more. In young 9-10- 
month-old males, sperm was formed a month later (N. Sokolov, 
1957). Decrease in the volume and loss of elasticity of the testes 
in arctic foxes on farms takes place at the end of April— beginning 
of May when the prostate gland ceases its activity. The male 
produces sperm during the course of 2—3 months. The testes of 
arctic foxes are larger in size compared to those of red foxes and 
the ejaculate contains a large number of spermatozoa. Among 
captives, the arctic foxes with the least weight in summer but 
which rapidly increased in autumn, were found to be the best re- 
producers. 

In the course of the winter, ovaries and oviducts of females in 
the pre-estrus condition develop earlier in adults than in young. 

Ovulation of the spontaneous type does not depend on the time 
of mating. Before the beginning of ovulation, the ovary of the arctic 
fox is larger than that of the red fox, and although the ova are 
smaller, the number of Graafian follicles is larger; this fact deter- 
mines the fertility of this species. 

Estrus in arctic foxes is more prolonged than in the red fox, and 
on animal sovkhoz [state farms], it lasts 12—14 days, but under 
natural conditions the female remains [in estrus] for only 4—5 days 

53Middendorf, 1869; Cherskii, 1919-1920; Paramonov, 1929; Barabash- 
Nikiforov, 1937a; Dubrovskii, 1940; A. Romanov, 1941; Rakhmanin, 1959; 
Skrobov, 1958; Chirkova et al., 1959; Starkov, 1932, 1937, 1940; Tavrovskii, 
1946; Il'ina, 1952 and others. 



342 

on the average. During estrus, the mammary glands enlarge to 0.5 
cm in diameter. In the copulatory period, the male and female pair; 
on farms this may last from one minute to several (4) hours (Boitsov, 
1937); coitus may occur repeatedly. 

In nature, the breeding season extends from the second half of 
winter to the end of summer. With the appearance of the first rays 
of the sun, courtship behavior begins. At this time, 1—2 males, rarely 
more, run after one female (in captivity, 2—3). The peak of rut and 
mating are often observed in March-beginning of April. Adult ani- 
mals commence breeding first, and then the young (on Yamal, 
10-15 days later), and last are those which have returned from 
migration. 

In various regions of the USSR, over a wide area the breeding 
dates of arctic foxes are nearly the same (Table 28), but the ap- 
pearance of young in the northern subzones of the tundra is normally 
2—3 weeks later compared to the southern (Yamal, Taimyr). 
239 In one or another place, the breeding season and fertility of the 
arctic fox can change annually in connection with the dynamics of 
nutritional conditions and age composition of the populations. The 
mating period in one place fluctuates within 2—3.5 months, and the 
time of birth of young arctic foxes is shifted by 1.5—2.5 months on 
the continent and up to 3 months in the Commander islands. Under 
favorable conditions, a local concerted mass rut is observed; in a 
year following inadequate food, it is undetectable, prolonged, or 
greatly delayed. In certain years, fertilization and implantation occur 
in 70-80% of the females and, in some places, possibly even more. 
At this time all littering burrows are occupied by young, and some 
arctic foxes deliver their newborn under clumps of grass. In other 
years, the majority of females remain barren and 10-15% or less of 
the littering burrows are occupied (see also p. 365ff.). 

After mating, arctic foxes explore the region which they have 
chosen, and the female begins clearing the burrows. This takes 
place at various times of day depending on the general course of 
reproduction, mainly in connection with the nutritional conditions of 
the year. In the Solovetsk islands, arctic foxes began to choose the 
place for the future nest but the burrows were ready 5-15 days 
before parturition. One female which was bred on 19 April, occu- 
pied a burrow together with a male on 2 1 May, and gave birth on 
6 June. In Yamal, the burrows are also cleared from March to the 
beginning of April — and on Belyi Island, to 20 April. In the lower 



343 



Yenisei, already in March, a female wandered around a hillock 
which she chose for her future lair, making a round of up to 5 km. 
In the northwestern Taimyr peninsula the clearing of the burrow 
progressed from the second half of April, and in Lena-Khatanga 
territory — ^the middle of March; a pregnant female littered here on 
7 June, and emerged after parturition on 10 June. On Mednyi 
island, the Arctic foxes now distribute themselves among the bur- 
rows after mating. 

In captivity, gestation continues 49-56 days, 52 on the average 
(also confirmed by observations in nature). In the earliest and latest 
parturition (45th, 62nd* day), newborn often die. The abdomen 
obviously increases in size 1—2 weeks before pupping, sometimes 
earlier. The female usually whelps in a burrow, but sometimes 
directly at the entrance: this is often a young female that did not 
prepare the nest, and later, are carrying the newborn to the burrow. 
The duration of parturition depends upon litter size, after approxi-. 
mately half an hour, the first pup appears; sometimes, parturition is 
extended over 2-3 days. Most often, the female delivers herself of 
her litter** at night or early in the morning. 

In the tundra, mass appearance of the newborn arctic foxes 
occurs in May and June and coincides with fawning of reindeer, the 
return of willow ptarmigan, and the arrival of geese, gulls and other 
birds. The earliest parturitions of arctic foxes known are mid- April 
on the Commander islands, the end of April on Novaya Zemlya and 
Belyi Island, and the very latest, in June in Sevemyi (North) Yamal, 
Lena-Khatanga territory. Commander islands, and other places. 



March 



April 




Fig. 89. Times of mating of arctic foxes in island husbandry on Kil'din, 
Murman (from Zolotov, 1940). 



*Appears inconsistent with previous values for length of gestation— Sci. Ed. 
**In Russian original, literally "burden" — Sci. Ed. 



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345 

The appearance of litters near burrows was observed from the 
end of May until August, and more often from the second half of 
June to the beginning of July; it is very rare when young appear 
later than August. The appearance of delayed litters in August 
("of a mitten" size) or when still in summer coat (krestovatik)* 
in November, is often inteфreted as a second litter: Actually, the 
late whelping is explainable as delayed estrus or a second estrus 
241 after failure to conceive, abortion, or death of the newborn. Cases 
of repeated estrus in captives are known, up to 4-5 times 7-30 days 
after the first mating. In some years, 5—8% of the females come 
into estrus for a second time. Apparently, the males, in some cases, 
retain the capacity to mate with the female in a later period. Thus, 
the testes of an arctic fox from Yakutiya, taken from its family at 
the end of July, still contained mature sperms (N. Sokolov, 1957). 

The emergence of the litters from the burrows takes place in 
the course of August— October, depending on nutritional conditions, 
and in the unfavorable years (Kara region), even in July. In years 
of abundant food, the litters remain near the burrows during the 
course of the entire autumn and winter. 

Fertility of the arctic fox is very high for a carnivore. Fertility 
can be judged by the number of corpora lutea in the ovaries, of 
embryos, and of the remains of the post-partum pigmented spots 
[placental scars] in the uterus. The latter are retained until the 
beginning of proliferation of the reproductive organs prior to the 
next estrus, and sometimes (in starvation) to the new estrus. In 
1934, when food was abundant on the Kolyma, intensive reproduc- 
tion was proceeding among the arctic foxes. In 27 females studied, 
the number of placental scars in the winter 1943/44 was from 3 to 
24 (in the majority — 10, 11, 15 and 17), averaging 12.7 for each 
parturient female (Cha, 1953). 

The situation becomes more complicated when the population 
decreases. Thus, in the Kara tundra, the summer of 1956 was not 
favorable for reproduction of arctic foxes due to the disappearance 
of lemmings. Out of 22 adult females investigated in the winter of 
1956/57, 8 remained barren (placental scars in uterus absent); 6 
females revealed from 1 to 6 ill-defined scars, and the remainder 
from 6 to 16 well-defined scars at the same time (in one even in 
the cervical region of the uterus) and up to 10 ill-defined patches 

*Local word; see p. 351 — Sci. Ed. 



346 

in each. The latter might have been traces of the placentae of 
embryos resorbed before parturition. The largest number of scars 
was 21. On the average in each pregnant female about 1/3 of the 
embryos did not live until parturition; on the average, of 10.2 em- 
bryos, only 7.1 were likely to have been bom (Chirkova et al., 1959). 
The greatest number of placental scars found in one arctic fox 
on Belyi Island was 26 (Tyulin, 1938). The possibility of one female 
nursing such a large number of pups has never been confirmed. 
Animal breeders report up to 16 newborn delivered by one mother. 
Cases of finding more than 20 young arctic foxes in one burrow are 
most frequently explained by the union of two or more litters. Thus, 
in the Kolymsk area, when a colony of burrows was excavated, up 
to 40 or more pups were found. On the Commander islands, a case 
was noted of two females with their pups of various ages living 
with a pair of parents with its own litter in one burrow. Here also, 
females were observed who, beside their own offspring, had 3-4 
foster pups. When breeding arctic foxes in captivity, a maximum 
number of 22 embryos was observed, but usually a part of them 
was resorbed or the pups were bom dead. 

Geographic variation in fertility is observed between continental 
and maritime arctic fox populations. In continental tundras and on 
Novaya Zemlya, great fluctuations are observed in litter size and 
the proportion of infertile (unmated) females in the various years. 
The number of pups in a litter in the continental tundras varies from 
8-12 to 20 and more^"* on the average in years with abundant food, 
and decreases to 3-5 in years of famine. In the maritime tundras, 
for example on the Kola peninsula, in northeastern Siberia, and 
especially on the Commander islands, the size of litters (to 10-13 
pups) and the number of females participating in reproduction 
varies to a lesser extent from year to year. On Kil'din island, poorly 
nourished females produced 4—6 pups, and females nourished better 
than average — 11-13. On the Commander islands, reduction was 
242 repeatedly observed in the number of young in litters fi^om 6—7 to 
2—3 on the average in separate places — ""ukhozh"* depending upon 
the abundance of natural food and supplemental food. The absence 
of large litters is also characteristic for the island husbandry in 
North America and was noted in the maritime populations of arctic 



^To 22 in Yamal (Rakhmanin, 1959). 

*Literally, "pasturage"; in this context, foraging places — Sci. Ed. 



347 

foxes in Greenland (Lek, 1957)*. Apparently the character of this 
variability is conditioned by the greater or lesser constancy of the 
supply of food. 

Therefore, the fertility of arctic foxes is lowered according to 
the number of infertile and unmated females as well as to the death 
of offspring — ^the resoфtion of litters during the time of embryonic 
development and the death of pups after birth. In fox farms, unmated 
female averaged 7% at the end of the 1930's and those which 
were infertile — 15%. Embryonic mortality attained 36%. 

The sex ratio in populations of the arctic fox is almost 1 : 1, but 
this ratio may change depending on season, food conditions of the 
year and the age composition of the population. In the majority of 
cases observed, males predominate. In Kola and Saltykov fur 
sovkhozes [state farms], 2—3% fewer females were bom. In the 
latter, during the period 1950-1957, deviations established a pre- 
dominance of males in one year in one sector of the sovkhoz up to 
4.4% (of 1 100 pups) and the dependence of sex [in litters] on the 
age of females; in the litters of young (up to 2 years) and the old 
(6-8 years) females, males predominated (5 1-54%), while in those 
of moderate-age — females (51-51.5%). An increased number of 
males was observed in large litters (Ivanter, 1962). 

On Yamal, at the beginning of the 1930's, 50.5% males were 
caught in one party, but here in 1933/34, females predominated (5 1 .6%). 
In Bol'shezemel'sk tundra, at the beginning of the winter of 1956/57, 
the take of males was 56.4%, but by the middle of winter, there was 
no difference. On the Taimyr peninsula, at the beginning of the winter 
of 1931/32, 64% females were caught. Among 2182 arctic foxes ob- 
tained on the Commander islands, males constituted 53.8%. 

Under natural conditions arctic foxes are usually monogamous, 
but individual cases of polygamy are known, it being observed on 
the Commander islands each year. This capacity of arctic foxes is 
used in breeding the animals on farms where only one male is kept 
with 4—5 females; individual males can accomplish 20-25 matings. 
Cases of polyandry, observed on the Commander islands, are ex- 
tremely rare. 

The ratio of young to adult animals in arctic fox populations 
changes considerably depending on feeding conditions, time of year, 
and on their total number. On the Lena, 62.9% young were caught 

*Not in Literature Cited.; probably Lack, 1954, p. 46 — Sci. Ed. 



348 

in the good year of 1935/36. In the Kara region in the season of 
1956/57, a quite abundant one, during the time of peak migration 
more than 2/3 of the females were young"; in the second half of 
winter, the latter constituted only 1/3 of the animals taken. In Yamal, 
the prevailing majority of individuals in the populations (in three 
harvest seasons from 1955/56 to 1957/58 inclusive; 475 individu- 
als — 94.7%), were determined as juveniles*. The ratio of adults to 
young animals constituted 1 : 18.9; this explains the selection of 
harvesters in direct ratio to the young animals (Smirnov, 1959, 
1959a). Such abundance of young animals over a duration of 3 
years otherwise seems improbable. 

Arctic foxes reproduce well for 1—2 years in succession, alter 
which they "rest" for 1-2 years. This periodicity observed 
repeatedly in the 1930's in the Kola fox sovkhoz was explained by 
"physiological exhaustion" of the female organism in which the 
first symptoms of overwork were revealed in the third year after 
the first two; the older the female, the lower the probability of its 
participation in reproduction at this age. Arctic foxes of late litters 
(in captivity) do not always reproduce the next year. 

By means of the artificial insemination of silver-black foxes 
with arctic fox sperm, the mating of female red fox with blue arctic 
fox, and blue arctic fox female with the silver-black and white- 
243 muzzled foxes, hybrids of arctic fox and red fox were obtained. 
These hybrids had the intermediate characteristics of both species 
but all of them were shown to be sterile. 

Growth, development, and molt. White arctic foxes are bom 
covered with dark smoky-brown short fur, while blue arctic foxes — 
with dark-brown, almost chestnut. As in young red foxes, newborn 
arctic foxes are born blind, without teeth, and with ear pinnae 
closed. They are smaller than [newborn] red foxes and weigh from 
60 to 85 g; their body length is 1 1—13 cm; tail, 4.2—6 cm. Soles of 
paws are bare, but already by the fourth day they begin to develop 
for. 

Growth in the animals proceeds more intensively than in young 
red foxes. Eyes open on the 9th-18th day, simultaneously with the 
opening of the external auditory meatus. Eye color darkens over 
time. In the blue arctic fox, the eyes are bluish-gray at an early 

^Uterine determination of age (Chirkova at al., 1959). 

*The Russian word sevoletki literally means "this summer's"— Sci. Ed. 



349 

age; they turn yellow in approximately the 7th month. On the 12th 
day, the irruption of the incisors can be noted, and in some pups, 
the camassial teeth also. On the 15th day all the incisors on both 
jaws have irrupted. Canines appear first in the lower, and then in 
the upper jaw. On the 18th day, the premolars are cut, first in the 
lower jaw. All milk teeth have appeared by the 27— 28th day. Re- 
placement of [deciduous] teeth by the permanent ones takes place 
in wild arctic foxes in the end of summer and the beginning of 
autumn in the cross and blue fox types (see below) (Zhitkov, 1913; 
Kozlova, 1933). 

In one-year-old animals on Yamal in winter, the teeth are snow 
white and bright without any sign of wear. In contrast to the red 
fox, tooth wear with age begins with the upper incisors, proceeding 
to the first and second molars of the lower jaw. Earlier than in red 
fox, the fourth upper premolar is worn down and, later than in this 
species [red fox], the first and second molars of the lower jaw [are 
worn down]. Age determination by tooth wear was shown to be 
possible by also taking into consideration the changes in the propor- 
tions and configuration of the skull (Grigor'ev and Popov, 1952). 
Age determination according to the degree of closure of the central 
canine cavity is also possible. In this case, only the first two years 
age groups can be clearly differentiated (Smimov, 1959, 1959a). 

With age, the skull undergoes changes. In the young continental 
arctic foxes still with milk teeth, the posterior region of the skull is 
higher, and the nasal region is shorter and wider than in adults. 
Orbits smaller. Zygomatic arches thinner and narrower, bullae osseae 
more swollen and rounded, hard palate wider, sagittal crest absent 
or weakly developed. Later, the skull broadens in the area of zy- 
goma and frontal region, which is abruptly elevated. The postorbital 
processes are large and the postorbital constriction is well defined. 
Muzzle elongates, sagittal crest is greatly enlarged, especially no- 
ticeable in Commander arctic foxes (Ognev, 1926). 

At the age of 5-7 years, all teeth in Commander arctic foxes 
are worn down. Duration of life of arctic foxes assigned to island 
husbandry and sovkhozes for breeding service is 6-10 years. A 
case is known of an arctic fox living in a zoo for 20 years, when 
the animal died in a state of complete senility (Paramonov, 1929; 
Boitsov, 1937; Il'ina, 1949-50). Under commercial conditions, arctic 
foxes very rarely survive to old age. In the forest-tundra on the 
Yenisei, a very old arctic fox was caught in which only the crooked 



350 

remains of completely obliterated teeth were left; the pelage was 
short — ^about 2 cm in height, tail thin, but the animal itself looked 
very large (A.F. Chirkova). On Yamal, in the winter of 1958/59, 
when the number of young was small, only 5.5% of the arctic foxes 
caught were older than 3 years of age (Smimov, 1960). 

Arctic foxes grow quickly, especially at first. Thus, the average 
daily weight gain on farms decreases fi^om 13% of body weight in 
the first days to 2.5% at the age of 2 months. Increase in weight 
depends also on food composition. On Kil'din Island husbandry on 
a mixed ration gave a daily weight increase of 20-40 g from days 
244 42 to 90, an average of 28 g; those kept on fish ration, gave an 
average daily weight increase of 4 1 g and those on seal meat — 54 
g. Increase in length changed also. In farms, the average weight of 
pups at the age of one month was equal to 600-650 g, at two 
months — 1.7-1.8 kg, at three — ^2.5 kg, at four — ^3.8 kg, at five — 4.5 kg 
and at six months— 5 kg. At 6-7 months, the young arctic foxes 
have almost attained adult size. From the age of 7 months to 2.5 
years the weight of arctic foxes on farms increases on average 
from 5.6 to 7.1 kg in males, and from 4.9 to 6.2 kg in females. 

The change in the weight of the pups of the early and late 
litters is greatly different: on a Kola fox sovkhoz, this difference 
reached 1.5 kg at the age of 4 months. As compared to the red fox, 
the growth constant of the arctic fox before sexual maturity is 
larger, and in the sexually mature animals, smaller. Growth of the 
arctic fox is more quickly completed than in red foxes (Boitsov, 
1937; Tavrovskii, 1946; Il'ina, 1952). 

In nature, young arctic foxes had the following dimensions: on 
Yamal, July 2, weight 1250 g, with body length 40 cm and tail 16 
cm; August 24 on the Khatanga, body length 51 cm and tail 21 cm; 
in October, on the Kolyma, a young blue fox (see below) weighed 
3 kg with body length 54 cm and tail 32 cm; the latter animal had 
already attained adult dimensions (Zhitkov, 1913; Tsetsevinskii, 1940; 
A.F. Chirkova). 

Seasonal variations are well manifested. Body weight of arctic 
foxes is lowest in summer and highest in December. In connection 
with rut and mating, weight drops greatly in the second half of 
winter, especially in males. On the Commander islands, it decreased 
by 15-20% as compared to that in December-January (Boitsov, 
1937; Il'ina, 1940, 1950). 



351 

In the white arctic fox, the following age and seasonal stages 
of the pelage are differentiated (Paromonov, 1929; Boitsov, 1937; 
Guber, 1939; Rakhmanin, 1948; N.N. Spitsyn). Slepushonka* [mole- 
vole] or kopanets* [digger] has a delicate dark-brown, almost black 
fur, with very sparse, separately protruding guard hairs. Skin length 
30-35 cm (without tail); age up to 1-2 months. In the nornik* 
[burrow dweller], the quantity of guard hairs is somewhat greater 
and the pelage turns browner; the back, shoulders, head and tail 
darken. Skin length without tail 40-45 cm. It still lives in the burrow; 
age 2-4 months. Krestovatik* [cross fox] — ^the arctic fox in sum- 
mer coat, distinguished by brightening of the pelage on the belly and 
along the sides, acquiring a grayish straw-colored tone. The brown 
stripe along the back and on the shoulders is clearly marked, in the 
form of a cross, and guard hairs are considerably better developed 
than in the preceding stages. Skin length of the young cross fox 45— 
60 cm, of tail — 30 cm. Sinyak* or chayachnik* [blue fox] has a 
shorter pelage; following upon the appearance of a large amount of 
white hairs, it becomes lighter and grayer, taking on a bluish, leaden 
tone. Skin length without tail — 54—68 cm, tail length — ^32 cm. Guard 
hairs up to 40 mm in length. Age, 6-8 months. Arctic foxes are 
found in this stage from the end of September to the middle of 
October. Replacement of the milk teeth by the permanent ones 
takes place in the krestovatik and sinyak stages. In the nedopesok* 
[premature], fur is grayish-white, with scattered brown guard hairs 
concentrated on the upper part of the body. Underfur azure-blue. 
The animal is found in this pelage from the middle to the end of 
October. Complete, full-grown or doshlyi* [finished] — this is the 
arctic fox in its winter white fur; some animals only retain a few 
"bead" guard hairs, and light bluish color at the base of the hairs. 
Wool is dense and fluffy. A few arctic foxes attain the full winter 
pelage by the end of November, but the majority in December- 
January. 

In full winter pelage, the so-called standard arctic fox is divided 
into 3 sorts depending upon the degree of maturity, whiteness, 
length and evenness of guard hair distribution, and the thickness of 
the entire pelage. Not all arctic foxes reach full fur development 
each year. The greater part of the skins prepared during the course 



*Translator's remark: These local names (on this and the following pages) are 
transliterated, and the literal English meanings given in brackets. 



352 

of a winter are of the second and third sorts. In first year arctic 
foxes, fill! development of fur cover is rarely attained. The most 
high quality furs are supplied by 3-4 summer [2-3 -year-old] males 

245 and 2-3 summer [1-2-year-old] females. Emaciated arctic foxes, 
and individuals, often frequenting burrows, or living within thickets 
of stunted tal 'nik [риф1е osier] are distinguished by dull and frayed 
fur. 

At the end of winter, when the still white and fluffy fur of the 
arctic fox begins to weaken and "leak", the animal enters the 
veshnyak [spring] stage. Later, usually in April, during the forma- 
tion of the summer hairs and the darkening of the flesh side of the 
skin, the almost guard-hair-less skin is called "gagara" [loon]. 
After shedding of the winter hairs, the adult arctic fox again enters 
the summer stage — krestovatik, which differs from the young ani- 
mal by its firmer pelage; length of guard hairs is about 1 .5 cm. This 
completes the cycle of seasonal pelage change. Throughout the 
course of their lives, the adult white arctic foxes annually go through 
these described stages of pelage replacement, except for 
slepushonka and nornik. 

The pelage of the blue arctic fox changes with age and to a 
lesser extent, with the seasons. From the almost black or dark 
brown {slepushonka), the fur of the adult arctic fox gradually 
lightens, and acquires a dark violet to light-beige color. With age, 
it is mainly the tone of the hair color that changes. At the end of 
August on the Commander islands, the majority of arctic foxes 
already have a normal "blue" colored winter pelage. Some fur 
categories are differentiated — ^gray or brown tones on Bering Island 
and chocolate on Mednyi Island. Hair softness also varies. Arctic 
fox fur becomes completely "vykhodnyi" [grown out] in the sec- 
ond half of December— January. Average length of contour hairs in 
the 1.5— 2-month-old arctic foxes is about 27 mm, on winter skins, 
more than 82 mm; in winter, hair thickness also increases. As a 
result of the warm and humid climate on the Commander islands 
and especially in connection with the delayed arrival of cold, the fiir 
of those arctic foxes is distinguished by coarser and shorter guard 
hairs and often by matted underfur (Tret'yakov, 1871; Barabash- 
Nikiforov, 1937; Boitsov, 1937; Il'ina, 1949, 1950). 

In adult arctic foxes molt takes place twice a year, in spring 
and autumn. On fanns, arctic foxes have a full winter fur cover 

246 from December to February. Spring molt proceeds from the end of 



353 



». ■■ J-- 



V* '►^t,** 







.л*'^^» 



Fig. 90. Blue fox in summer fur. Mednyi Island. Photograph by 

N.N. Kartashev. 



February to June. Summer hairs are initiated in April and grow until 
the middle of August. Autumn molt occurs in August— September, 
while the growth and maturation of the winter pelage takes place 
in October— December. Guard hairs in winter are twice as long as 
in summer (an average of 45.8 mm, and underfur 37.9 mm), and 
the hair thickness is also greater. Underfur quantity constitutes 97% 
(Eremeeva, 1956; Kaletina, 1956; Kaletina et al., 1957). In winter 
there occurs an abundant deposit of subcutaneous fat. Sweat glands 
at this time are weakly developed. In summer, the fat layer is 
reduced and the size of the sweat glands increases. 

The beginning of spring molt coincides with the start of estrus. 
In the first instance, pregnant females molt, their skins losing winter 
fur just after parturition. Males, barren females and yearlings molt 
later, and the last are old and diseased animals (A. Romanov, 1941 
and others). Holding arctic foxes at higher temperatures (18 ± 20°C) 
and lower humidity (42 ± 8%) speeded up the time of spring molt 
by 8 weeks (Il'ina, 1926). 



354 

In the wild, the time of pelage replacement of arctic foxes 
varies greatly depending upon meteorological conditions, the abun- 
dance of food and nutritional state of the animals, their age, sex and 
the physiological condition of different individuals. Spring molt 
begins earlier and occurs more rapidly in years with good food 
conditions. In the southern tundra zone, spring molt begins 1 .5—2 
weeks earlier and ends 2-3 weeks earlier compared to the northern 
coast and the Arctic Ocean islands (Rakhmanin, 1959). Molt of 
arctic foxes in February was observed in different years on the 
Kola peninsula, Yamal and even on Belyi Island. More often, the 
onset of molt is observed in March, and on Taimyr and Novaya 
Zemlya, even in April. In various populations, it extends for 4 or 
more months and in the first 2 months is usually not obvious. Shed- 
ding of winter fur ends in the majority of tundras in June or July, 
but significant individual deviations are observed. On Klimets Island 
(Lake Onega) in the middle of June, a half-white arctic fox was 
killed (Sluchevskii, 1886). Individual unmolted arctic foxes are en- 
countered even in August. On Novaya Zemlya in the beginning of 
May and on Yamal at the end of May, some individuals were found 
in winter pelage, and on the New Siberian islands, arctic foxes 
were white even at the beginning of July (Krivosheya, 1884; 
Paramonov, 1929; Dubrovskii, 1937, 1940; Tyulin, 1938; A. Romanov, 
1941; Rakhmanin, 1959). In Greenland (Seton-Thompson, 1909- 
1925), arctic foxes are met with which, as in hares, are not able to 
change the winter pelage during the short summer and remain 
white throughout the year. 

Spring molt starts with the shedding of the guard hairs, then 
underfur which is, however, more quickly replaced than guard hairs. 
Replacement proceeds from the end of the muzzle, rump and tail 
base, spreading along the back in the shoulder region, on the limbs 
and later — on the side and belly. Winter hairs are retained longer on 
the tail. During the formation of the summer hair cover, darkening 
of the flesh side of the skin begins around the eyes, at the base of 
the tail, on the rump, forefeet and then spreads to the thighs and back. 

Autumn replacement of hair cover proceeds in the wild from 
September to December. Better fur occurs in January— February. 
Time of autumn molt is delayed in the south for 2—3 weeks in 
comparison to northern regions (Rakhmanin, 1948, 1959). Thus, 
earlier maturation of fur was observed on Belyi Island (about 
73° N. Lat.), where at the end of September 1930, arctic foxes 



355 

wandering over the snowless land were 8Ьаф1у distinguished by 
their already whitened skins. On the eastern coast of Sevemyi 
Island of Novaya Zemlya (Sedova gulf, 75° N. Lat., Ledyanaya 
Gavan'), during observations on 12 September and October 1936, 
arctic foxes in white coats were also encountered (Rundan, 1936; 
A.F. Chirkova). Fur maturation proceeds asynchronously in arctic 
foxes of various [fur] types. In Kara region, in December 1956, 
247 among arctic foxes of the Pechora type caught, 33.5% of the skins 
were of sort 1 [see above], and among arctic foxes of Obdorsk 
type, coming from the higher latitude of Yamal — 79.7% (Chirkova 
et al., 1959). On the Commander islands, maturation of fur at 
different times by habitats ("ukhozh")* is well known; animals 
from rivers in montane barrens have better coats. From the second 
quarter of this century, when the climate of the islands warmed 
noticeably, autumn pelage appeared later. Earlier, harvest began 
here in November— December, when the ftir of the majority of arc- 
tic foxes was already mature; at the present time, for gathering 
better skins, arctic foxes are killed at the end of January and in 
February (Il'ina, 1949-1950). 

Falling temperature and increased humidity, especially in Octo- 
ber, have a favorable effect. Humidity influences increase in guard 
hair length and fluffmess. Nutritional status, sex, age, and physi- 
ological condition of the individuals affect the time of autumn molt. 
It was noted that the more abundant the food in autumn and begin- 
ning of winter (but not in summer), the longer the delay in maturation 
of fur. Fur matures earlier in males, and later in barren females. 
The young of late litters develop full coat later than adults. Molt of 
old and diseased individuals is delayed. 

Full pelage growth takes place at different times in different 
years. On Yamal in the winters of 1923/24 and 1924/25, when the 
numbers of arctic fox was low, sinyak [see above] constituted 1.2— 
2.0%, i.e., winter fur of arctic foxes developed comparatively early, 
but in 1925/26, when arctic foxes were abundant, this group con- 
stituted 20%, nedopesok constituted 1 8%, and top quality fur was 
only 30%; i.e., growth of winter fur was greatly delayed. In the 
winter of 1927/28, when arctic foxes were few, nedopesok con- 
stituted 8%, and top quality furs were 45% (N.N. Spitsyn); i.e., 
growth of winter fur proceeded more intensely. In the lower Lena, 

*See previous footnote, p. 346 — Sci. Ed. 



356 

in the winter of 1935/36, when arctic foxes abounded, sinyak were 
encountered throughout the winter (in February 0.4%). Skins of 
sort 1 in the March collections were 23.2% — development took 
place slowly (Tavrovskii, 1939). In the Taimyr National Region in 
the winter of 1945/46, in all, 5.3% sort 1 skins were processed 
while in the winter of 1936/37 — 30.6% (N.N. Kuznetsov). On 
Novaya Zemlya, individual arctic foxes with dark ears and tail tips 
are sometimes seen throughout the winter. Arctic foxes with in- 
completely developed winter fur in any case begin spring molt also 
in February and March (Tsetsevinskii, 1940). Therefore, in years of 
comparatively good food conditions for the young, in years of a low 
number of animals, the development of the winter coat is acceler- 
ated, while in the absence of food in years of intensive reproduction 
of arctic foxes, it is delayed. 

For Lena-Khatanga territory the following scheme was pro- 
posed for seasonal changes in hair cover of the arctic fox (A. 
Romanov, 1941)^^ 

January— February — ^all arctic foxes "doshfyr [finished], skins 
sorts I, II and III. 

March — predominance of first sort skins; in second half of month, 
rarely veshnyak [spring] (the first molt stage). 

April — ''veshnyak" predominates; in second half of month 
''gagara" [loon] appears (beginning of intensive molt). 

May — ""gagard" predominates (intensive molt). 

June — all arctic foxes in ''gagard" stage (end of molt). 

July — all arctic foxes transit to krestovatik [cross] stage. 

August — all arctic foxes in krestovatik stage. 

September — krestovatik predominates, a few sinyak [blue], 
rarely, nedopesok [premature]. 

October — krestovatik, sinyak, nedopesok; rarely ""vykhodnyi" 
[grown out]. 

November — sinyak, nedopesok; vykhodnyi predominates with 
skins of lower sorts [II, III]; a few first quality. 

December- — vykhodnyi skins of all sorts, sort II predominates; 
nedopesok is rare. 



^Other evidence of A. Romanov was also taken into consideration on the 
development of arctic fox fur in the winters of 1925/26, 1926/27 and 1934/35, with 
different environmental conditions. 



357 

The topography of hair cover replacement in autumn differs 
from that in spring. Molt also begins with the head, which turns 
gray on the sides, but subsequently, whitening takes place on the 
groin and on the sides of the body, and later on the back and tail. 
On the back, individual dark hairs are retained for a long time. 
248 Sometimes, whole areas of the fur on the nape remain unmolted. 
During the time of the autumn molt the flesh side of the skin in 
white arctic foxes remains light, but in the blue, it darkens; at first, 
the flesh side grows thick, but later becomes thin. 

Among the pelage variants of white arctic foxes, cases of 
partial chromism in the form of 8Ьаф1у delimited spots of foxy color 
are noted. In the Bol'shezemel'sk tundra, white arctic foxes were 
caught with spots of reddish fur on the side or on the posterior part 
of the back, from 2—3 to 10—15 cm and larger. Spotted individuals 
were observed in the years with an abundance of arctic foxes. In 
January 1947, in Nizhnii Kolymsk region, an arctic fox was caught 
with a bright-reddish spot 6.3 x 19 cm in diameter. 

Enemies, diseases, parasites, mortality, and competitors. 
Enemies of the arctic fox are few. Among them are the wolverine 
and wolf, which are found in tundra in some numbers. The polar 
bear rarely attacks the arctic fox, but in the years of famine, it may 
harass the fox to death. The main enemies are sled and herders' 
dogs. Among birds, the golden and white-tailed eagles, the large 
species of gulls (glaucous and others) and skuas, raven and even 
crow are dangerous (to young), and on the Solovets islands, also 
goshawk. More often, the white polar [snowy] owl annoys the 
arctic fox, being especially numerous in the years of intensive re- 
production of the animal; in the taiga zone — eagle owls. Owls attack 
young arctic foxes during their dispersal from burrows and also kill 
those weakened by disease or migration. Owls are periodically very 
abundant in the tundra. Thus, in Bol'shezemel'sk tundra 600 birds 
were trapped in spring, summer and autumn 1955 in an area of 
about 400 km^ (Yu.V. Rybalkin). 

In years of insufficient food with an abundance of animals and 
intensive migration, epizootics are common in the migrants (Chirkova, 
1951, 1955; Sokolov and Cha, 1957; Meteleva and Rubanchik, 1959; 
Chirkova et al., 1959). Native inhabitants have long been acquainted 
with the disease called here dikovan\ or frenzy [rabies], "plague". 

Losing their sense of fear, furious arctic foxes run into villages, 
enter dwellings, attack dogs, reindeer and humans. Animals bitten 



358 

by arctic foxes become rabid, but the disease is extremely rare 
among people. Scars and gashes on the muzzle and other body 
parts of the arctic fox indicate that the animals often bite each 
other, and this influences the rapid spread of the disease. Usually 
many other species are involved in a rabies epizootic: red foxes, 
wolves, wolverines, ermines, weasels, sled and hunting dogs, hoofed 
animals and even birds — owls, gulls and others which feed on lem- 
mings. The disease among arctic foxes is usually preceded by the 
massive death of lemmings. 

The bodies of dead arctic foxes are not often observed in the 
winter period, and only after the disappearance of snow is the size 
of the epizootic revealed. Thus, in the spring of 1933, in the middle 
of Yamal, the remains of 16 arctic foxes which had died in autumn 
and winter were found along an extent of 20 km^ In the same 
year, in the Se-Yaga region, 130 dead bodies of arctic foxes were 
discovered in an area of 3000 km^; the majority were in winter fiir 
and high nutritional status. In the Pechoro-Ilychsk preserve, in the 
second half of winter after the massive migrations of arctic foxes 
in 1929/30 and 1935/36, disease was observed with the death by 
spring of nearly all immigrants. In the next season, disease was 
observed among dogs similar to that of arctic foxes and much 
mortality among them (Teplov, 1960; A.F. Chirkova). 

Large epizootics among arctic foxes took place, from the end 
of 1946 and in 1947 in almost all expanses of tundra in the USSR 
after the disappearance of lemmings (following unfavorable meteo- 
rological conditions in 1946 and 1947). By the spring of 1948, 
surviving arctic foxes were scarce everywhere, reproduction pro- 
ceeded slowly, and their numbers shaфly decreased by the season 
of 1948/49 in all tundras of the Soviet Union, except for the Far 
East where their reduction in number had occurred in 1947 
(Chirkova, 1955). 

At the beginning of the winter of 1953/54, in northern Yakutiya, 
soon after an outbreak of lemmings, disease appeared among arctic 
249 foxes. The diseased or dead constituted about 15% of those ob- 
tained, and all were well nourished — the arctic fox fed on the dead 
lemmings. The disease followed a swift course, and was transmit- 
ted to dogs, wolves and reindeer. Later, starvation of arctic foxes, 
migration and sharp suppression of reproduction were observed in 
1954; about 1/3 of the females were left unmated and more than 
half of them remained barren (Tavrovskii, 1958). Only among 



359 

arctic foxes of the Commander islands were epizootics not ob- 
served. The number of animals is always shaфly lowered as a 
result of epizootics, and the intensity of reproduction of the surviv- 
ing animals sharply decreases. 

The study of rabies — a neuroviral disease of arctic foxes — was 
promoted in the extreme northeast of Bol'shezemel'sk tundra from 
1954 to 1962 (Kantorovich, 1956, 1957, 1963). It was established 
that outbreaks of the epizootic among arctic foxes took place in 
1947/48, 1951/52, 1954/55 and 1955/56, with moderate disease in- 
cidence (up to 10 cases) in 1949/50, 1952/53, and 1958/59 and 
1961/62, with simultaneous cases among dogs and individual occur- 
rences among wolves, red foxes and reindeer. The disease was 
observed exclusively in the cold months of the year from Novem- 
ber to March. Examination of the brains of 1596 healthy arctic 
foxes over the course of 6 summers revealed that in general 30% 
were carriers; in years of massive outbreaks or moderate inci- 
dence — ^30-75%, and in the absence of the disease — ^3-6%. The virus 
was also isolated from red foxes. In healthy wolves, ermine, mouse- 
like rodents (Siberian lemmings, voles), owls and dogs, the virus 
was not detected, and therefore the arctic foxes were considered 
the permanent carriers of rabies infection in the far north. The virus 
was isolated from young arctic foxes, and from males twice as 
often as from adults, or from females. The virus was also isolated 
injhe parotoid and submaxillary glands in the years of outbreaks in 
20-30% of cases, but outside the disease period, in low numbers of 
arctic foxes, and in the absence of migration — only in individual 
cases. 

Rabies in arctic foxes was also studied in the tundras of the 
Nizhne-Kolymsk region during the epizootic of 1958/59. It was 
shown that in intramuscular infection of 6 arctic foxes from arctic 
foxes, red foxes and dogs spontaneously ill with rabies, the incuba- 
tion period ranged from 10 to 18 days, after which all arctic foxes 
became sick and died within 2-3 days. At that time hydrophobia 
disappeared. All animals died in a condition of outstanding good 
nutrition. Pathological changes in the diseased animals were re- 
vealed only in the form of hyperemia in the brain. Babesh-Negri 
bodies were not detected in the diseased arctic foxes examined. 
The retention of the active virus in the frozen brain was confirmed 
after 202 days. Not a single case of disease among people was 
established, even after being bitten by rabid animals. However, the 



360 

question of virus pathogenicity to humans through various routes 
and dosages was not studied (Strogov, 1961). The disease is char- 
acterized as a viral arctic encephalitis of animals (Stroganov, 1964). 

Under natural conditions, the arctic fox is susceptible to 
leptospirosis (infectious zheltukha), especially at a young age, but 
in a mild form. The role of agent of leptospirosis in arctic foxes, red 
foxes and other animals is served by Leptospira icteroanaemiae, 
and in addition, this disease in both species is induced by L. 
icterohaemorrhagiae (causative agent of Vasil'ev-Wel's disease 
in humans). Animals and humans are both susceptible to leptospirosis 
(Vyshelesskii, 1954). 

Outbreaks of epizootics among arctic foxes was observed in 
Bol'shezemel'sk tundra in 1909/10, 1926/27, 1929, 1932/33, 1937/ 
38, 1943, 1947/48, 1955/56 and 1957/58; on Novaya Zemlya, in 
1891, and also in 1927 and 1932/33; on Yamal in 1923, 1928, 1930, 
1932/33, 1934/35, 1935/36, 1947/48, and 1957/58; in northern 
Krasnoyarsk territory in 1915/16, 1920, 1923/24, 1926/27, 1935/36 
and 1947/48; in Lena-Khatanga territory in 1926/27; in northern 
Yakutiya in 1944, 1946/47, 1950/51, 1953/54 (from the Yana to the 
Kolyma) and in 1947/48 (Olenek-Lena); on the Anadyr in the be- 
ginning of the 1880's, in 1890, 1903/04, and on the coast in 1944/ 
45. 
250 On fur farms and nurseries [sic], arctic foxes suffer from 
paratyphoid which carries off to 50% of the young, especially of the 
late litters; this disease has been known in the USSR since 1933 
and is caused by the bacterium Salmonella cholerae suis and S. 
typhi murium. There are references to the presence of canine 
distemper, Aujeszky's disease, pneumonia and tuberculosis and rarely 
anthrax among arctic foxes. In years of outbreaks of the latter 
among reindeer, much mortality was also observed among arctic 
foxes (Lyubashenko, 1941; Vyshelesskii, 1954; Sludskii, 1954; 
Lyubashenko et al., 1957; Pankov et al., 1957; N.N. Spitsyn). 

Skin diseases of arctic foxes in the wild have not been studied. 
In years of this epizootic disease, arctic foxes appear with incom- 
pletely developed coats, with bald patches and other damage 
(Rutilevskii, 1939; Rakhmanin, 1948; Chirkova, 1955 and others). In 
the European north, in 1945/46 and 1946/47, bald patches and 
"stunted" pelage were observed in arctic foxes. At the mouth of 
the Pechora, in February, 1956, at a time of epizootic, an arctic fox 
with scabs and almost bald was caught. In tundras of the Far East, 



361 

in 1944/45, among a great number of sick arctic foxes, up to 70% 
"spoiled" coats were encountered. 

In two arctic foxes from Yamal in 1957 and 1959, 5 species of 
external parasites were found: the gamazid mites Laelaps lemmi 
and Hirsutionyssus isabellinus, the fleas Ceratophyllus penicilliger 
and Leptopsylla ostsibirica and the biting louse of the snowy owl, 
Strigiphyllus ceblebrachys (Luzhkov, 1963). 

In arctic foxes of the Commander islands, scabies was ob- 
served in some cases, caused by Sacoptes scabiei (0.4%). On 
Bering Island, the mange mite (Demodex canis) was observed — a 
serious, usually chronic skin disease (7% animals infected), which 
was widespread among arctic foxes in the 80's of the past century 
and which was, probably, transmitted from dogs. On the Com- 
mander islands, the ear tick Otodectes cynotis and the biting louse 
Trichodectes canis were found on arctic foxes; sucking lice were 
observed [pediculosis] (1.6%) and they were struck with ringworm 
[trichophytosis] and moist ringworm (0.7%) infections. In 1935/36, 
of 61 arctic foxes examined, 27 were observed with these diseases 
and in 34 arctic foxes the biting louse and sucking louse were 
found. 

Infection with endoparasites among arctic foxes is almost 100% 
(Isaichikov, 1931; Dubrovskii, 1937, 1940; Kiфichnikov, 1937; 
Kolmakov, 1937; L. Popov, 1939; Rutilevskii, 1939; Shastin, 1939; 
Afanas'ev 1941; Petrov, 1941, 1958; A. Romanov, 1941; Pereleshin, 
1943; irina, 1949, 1950; Dubnitskii, 1953, 1957; Chirkova et al., 
1958, 1959; Britov, 1959; Petrov and Kosupko, 1959; Ovsyukova, 
1961; Luzhkov, 1963). In the USSR, more than 30^^ helminth spe- 
cies were found in arctic foxes, of Nematoda (1 1 species), Cestoda 
(10 species) and Trematoda (7 species) and two species of 
Acanthocephala (found in arctic foxes on fur farms). 

The nematode Toxascaris leonina is found in the intestine and 
stomach of arctic foxes nearly everywhere, and in the majority of 
animals; of 59 arctic foxes from the neighborhood of Kara, 84.8% 
were infected, on Yamal, of 118 animals in a year of depressed 
numbers of arctic foxes — 92.4%. On the Commander islands, the 
fluke Maritrema sp. (80—100% infection) and the cestode 
Mesocestoides lineatus (88% on Bering Island) are found; each 



^''There are 55 species of parasites known; one species of Protozoa, 44 
species of parasitic worms and 10 species of external parasites (Luzhkov, 1963). 



362 

individual contained some thousands of both species. The nematode 
Uncinaria stenocephala is very often met with, and infects young 
animals more intensively. The nematodes Spirocerca lupi and S. 
arctica invade the stomach (goose egg-size tumors); the latter 
induce extreme emaciation and susceptibility to rabies. Trichinosis 
is known in 34% of some populations. 

A great danger presented by arctic foxes is their infection with 
251 the cestode Alveococcus multilocularis, found in the adult stage in 
the intestine of 50.8% (among 59) of arctic foxes in the winter of 
1956/57 in the northeastern Bol'shezemel'sk tundra; in some ani- 
mals up to 200 thousand specimens were found. On Yamal, the 
infection rate in 1957-1959 reached 73.7% (in 118 arctic foxes, 
from 11 to 74 thousand specimens were found in each animal). In 
the Anadyr and lul'tin tundras (Chukotka), the total infection was 
62% (among 35 arctic foxes) with intensive invasion by hundreds 
and thousands of specimens (Ovsyukova, 1961). Apparently, it is 
this form of echinococcus that is found in arctic foxes on Kil'din 
Island, the Commander islands, and in continental arctic foxes of 
Yakutiya. This parasite, when present as large infection in arctic 
foxes, induces extreme emaciation and paralysis. On Yamal, arctic 
foxes are also heavily infected with the tapeworm Taenia 
polyacantha (in 86,4%) of animals) and T. crassiceps (in 46.6%). 
Trichinella spiralis, which induces a rise of temperature and pa- 
ralysis of respiration resuhing in death or chronic illness, was 
found in arctic foxes of the Commander islands and in some other 
regions of the Far East (Petrov, 1941). 

Species of massive infection — echinococcus and the majority of 
taeniae — ^are transmitted through small rodents. The burrows of arctic 
foxes are the foci of toxascaridosis. Echinococcus as well as the 
trematodes Opistorchis felineus and Metagonimus Jokogawai, 
the cestodes Diphyllobothrium latum, D. erinacei and D. caninum, 
and the nematodes Trichinella spiralis and Toxascaris leonina 
are pathogenic to humans also. 

The majority of helminth species of arctic foxes are found in 
the gastrointestinal tract, mainly in the small intestine (in 100% on 
Yamal); one trematode species— in the bile ducts of the liver, 1 
nematode — ^in the urinary bladder, 2 nematodes — in the bronchi, tra- 
chea and nasal cavity, and 1 nematode — in the stomach. The larval 
stage of one species of cestode lives in subcutaneous cellular tis- 
sue, and one trichinellan — in the muscles. Coccidians were found in 



363 

the liver and intestinal tract of the arctic fox. Each arctic fox 
(Bol'shezemel'sk tundra, Yamal) is infected by up to 4 species of 
endoparasites. The degree of parasitic infection on the Commander 
islands is related to the nutritional condition of the animals 
(Afanas'ev, 1941). In almost all arctic foxes, enteritis is noted as 
a result of the effect of helminths; in the presence of heavy infec- 
tion with alveococci, there is subacute necrotic enteritis. Helminthiasis 
reduces fertility and induces the death of animals. Helminths of 
arctic foxes are shared in common with red foxes, wolves (4 spe- 
cies on Yamal) and dogs. Infections of arctic foxes in the northern 
regions are less than in the south (L. Popov, 1939; Rutilevskii, 1939; 
Chirkova et al., 1959). 

Death of arctic foxes due to basic hardship frequently takes 
place in the cold period of the year. During ice-glazed periods, the 
arctic fox's claws become broken and it cannot obtain food in the 
littoral zone and the tundra. Arctic foxes caught on the thin autumn 
ice of the open sea often perish in great numbers. Along the west- 
em coast of Yamal and in Baidaratsk bay, dead arctic foxes are 
collected in tens or hundred in some years. Many animals die at the 
time of seasonal migrations in the unusual situations of the forest 
zone and Arctic seas, whence they come in hundreds in autumn, 
and in spring return individually. Arctic foxes often drown crossing 
large rivers, for example, the lower Pechora with its numerous 
branches. On the Nizhnii Taimyr, of 46 arctic foxes swimming 
across, an observer saw three drift out to sea and three die in the 
river. 

Several facts attest to the great vitality of arctic foxes. On the 
lower Lena, an arctic fox was noticed with a badly broken paw; 
gangrene had already appeared, and it weighed only 1075 g. On the 
Commander islands, a blind female with cataracts in both eyes, 
stumbling against rocks while running, brought into the light a litter 
and nourished them. 

On the Commander islands, arctic foxes died from eating salt 
fish. Dead animals were found with pathological changes in the 
252 kidney and with ulceration of the stomach and intestine; cases of 
pneumonia and emphysema were noted. Sometimes inexperienced 
mothers bite, and tear at, or drag about their young. Deficiency of 
milk may incite such mothers. Newborn young die mainly in the 
first days after parturition. Mortality among pups up to 10 days old 
constituted 22-24%, from 10 to 30 days old— 3%, from 1 to 2 months— 



364 

8%; i.e. the total number of deaths among young at the end of 
lactation was 34% (of those bom). From 2 to 5 months of age, 3% 
of the whelps died, and total mortality through the period of attain- 
ment of sexual maturity — ^about 40%. On the Commander islands 
mortality through yearling age of 33% was noted; on farms in 
Canada — ^33-46%. In the Commander island enterprises in the 1930's, 
mortality among arctic foxes in July— August around an "ukhozh" 
[foraging site] varied from 8 to 15%. Mortality among adults con- 
stituted about 5% per year. 

On Mednyi Island, of arctic foxes that died in the period of 
1930-1935, 20% died from injuries and fights, 13%) from poisoning 
by salted products, about 37% from diseases and 30% from other 
causes. On Bering Island, 21%» of the arctic foxes taken in 1937/ 
38 were diseased. 

Under natural conditions of the continental tundras, the magni- 
tude of mortality, like that of reproduction, varies considerably in 
various years as a result of the great fluctuation, above all in the 
food supply. Judging by post-parturition placental scars in the uteri 
of arctic foxes in northeastern Bol'shezemel'sk tundra in the sum- 
mer of 1956 (a year with decreased number of arctic foxes), an 
average of 3.1% embryos were resorbed per pregnant female; 50% 
of the young females had died by the beginning of harvest. On 
Yamal in the summer of 1933, with negligible numbers of lemmings, 
about 80%) of the young died. Almost a two-fold reduction in the 
number of young at the burrows was noted by "Sluzhboi Urozhaya 
[Harvest Management]" from May to September in various tundra 
regions of the Far North. To control mortality of young arctic foxes 
in the wild, local kolkhoz [collectives] and sovkhoz [state farms] 
bring food to the burrows in food-deficient years in spring and 
summer, as well as in autumn in places of approach and harvest of 
the animals (Guber, 1939; Tsetsevinskii, 1940; G. Naumov, 1941; 
Skrobov, 1958; Chirkova et al., 1959). 

During the past decade, a northward shift in the southern limit 
of reproduction has been observed. A series of authors (Rozanov 
et al., 1932; Skrobov, 1958, 1958a, 1960; Uspenski, 1963) consider 
this to be due to the colonization of red foxes in the forest-tundra 
zone, and to its deep penetration into the southern zone of tundra. 
However, the importance of the red fox as a competitor of the 
arctic fox is exaggerated, since the main food of the red fox is gray 
voles [Microtus] and of arctic fox — lemmings and marine flotsam. 



365 

Arctic foxes prefer to den in open, elevated parts of the tundra, 
while for the red fox it is river floodplains, mainly forested sections. 
The number of red foxes in the zone of contact of the ranges of 
both species is extremely small; in northern Arkhangel'sk district, 
in preparing skins during 28 years (until the season of 1962/63) for 
one red fox there were more than 10 arctic foxes and the differ- 
ence between the number of both species increased in recent years. 
The process of displacement of the limits of reproduction of both 
species is governed, one may suppose, by changes in a series of 
abiotic and biotic factors as a result of Arctic warming; i.e., the 
effect of warmth on the soil and its moisture, on the microclimate 
of the burrow, on the duration of snow cover, on the time of the 
vegetative growth period, perhaps on the displacement of the limits 
of some plant and animal forms, in particular, the distribution of 
species of small rodents — lemmings, which are the main food of the 
arctic fox and gray voles, preferred by the red fox. Attacks on 
young arctic foxes by red foxes were observed, as well as of arctic 
on red foxes, in the course of ousting of one or the other species 
from burrows and places of activity (Rozanov et al., 1932; Boitsov, 
1937; Dubrovskii, 1940; Skrobov, 1958, 1958a). 

Direct competitors of the arctic fox for food are the ermine and 
weasel, but because of the small number of these species in the 
tundras, they do not inflict large losses on the arctic fox. In the 
lower Khatanga, where the arctic fox appears in great numbers 
253 there, the ermine disappears. In years of lemming abundance, they 
are also eaten by wolf, polar bear, wolverine, wild and domestic 
reindeer and even shrews, and thus they compete with the arctic 
fox. On Kil'din island, contests with river otters for fish were 
observed, and in the littoral zone of the Commander islands — ^with 
swines for marine flotsam. Sometimes, the wolverine uses burrows 
of the arctic fox. In its turn, in Yakutiya the arctic fox occupies the 
empty burrows of the tarbagan marmot and in the mounds of ground 
squirrels. 

Everywhere in the tundra, the snowy owl competes with arctic 
fox for its main food which is also the small rodent; in years with 
many rodents, both the snowy owl and the arctic fox reproduce in 
great numbers in the tundra. The same was observed for the rough- 
legged hawk [Buteo lagopus], which is less numerous in the tundra. 
Small rodents are also eaten by 2-3 species of skuas [Stercorarius], 
peregrine, merlin, in some places short-eared owl, white-tailed eagle. 



366 

goshawk and, in addition, raven, crow, magpie and gulls; gulls and 
corvids feed together with the arctic fox in the littoral zone 
(Osmolovskaya, 1948). 

Population dynamics. Seasonal dynamics in numbers of the 
arctic fox in years when its main food is plentiful is well mani- 
fested. Minimal population density is observed in spring, and the 
maximum at the end of summer-beginning of autumn, just after the 
litters abandon the burrows. When food is scarce, the growth of the 
population in summer is sometimes so negligible that the number of 
arctic foxes in autumn is even less than that in spring. 

Annual variations in their numbers are extraordinarily sharp. 
Beginning in the middle of the last century, a 3-year repetition of 
"productive" years for the arctic fox was often noted, with fluc- 
tuations of from 2 to 6 years between years of abundance (Vrangel', 
1841; Brandt, 1856; Middendorf, 1869; Nosilov, 1909; Folitarek and 
Chirkova, 1930; Burke, 1936; Dubrovskii, 1937, 1939, 1940; 
Kiфichnikov, 1937; Guber, 1939; Rakhmanin, 1948; Chirkova, 1955; 
Sdobnikov, 1958 and others). During 35 years at the end of the past 
century and the beginning of the current one, the maximum supply 
of arctic fox skins at Irbitsk fair was observed each 3-5 years, and 
at Yakutsk each 1-5 years (Silant'ev, 1898; Paramonov, 1929). 
During 1924—1959, maxima of prepared skins were repeated each 
2-4 years^l In separate regions of the north, after the October 
Revolution, the interval was 1^ years (2-3 years on the average). 
During the last decade, some stabilizing of fluctuations is observed 
in the arctic fox catch. Apparently, this is a result of technical 
measures to improve hunting. The average catch of arctic foxes in 
this period increased (see p. 376). Fluctuations in the catch and in 
preparation of skins correspond, in general, to the dynamics in the 
number of the species revealed by the more or less accurate count 
of wild animals in nature. 

There is great amplitude of variation in the numbers of arctic 
foxes in successive years. In the autumn of 1912, in the vicinity of 
Russkii list' [Russian Mouth] of the Verkhoyansk region there was 
an extraordinary "raid" of arctic fox, "what mosquitos" as ex- 
pressed by the natives (Zenzinov, 1914). Following such a year of 



^^Not counting the war years, when in the USSR as a whole and in Krasnoyarsk 
territory, 5 years passed between 2 successive peaks, probably as a result of 
special economic conditions. 



367 

abundance, the arctic fox was rarely met with. The number of 
animals in census areas varies tenfold in various years. The number 
of breeding burrows changes from a small percentage in unfavor- 
able years (Table 29) to frill occupation of all breeding burrows and 
the appearance of pups even in temporary shelters in years of 
abundant reproduction of the species. In 1935, in the Kara region — 
in northeastern Bol'shezemel'sk tundra, in a year of growing 
numbers of the species, 39% of the burrows found were occupied 
by large litters and the total number of arctic foxes on the experi- 
mental plots increased more than 9 times as compared to the 
254 preceding year, when only 5% of the burrows were occupied and 
litters were fewer. In the same region in the winter of 1957/58, as 
compared to the preceding winter, the number of arctic foxes de- 
creased 3.7 times (N. Dement'ev, 1955; Chirkova et al., 1959). On 
the northern shore of Taimyr Lake and along the Nizhnii Taimyr 
river, with intensive reproduction of arctic foxes in 1947, about 70% 
of the old burrows were occupied; in 1948, with weak reproduction, 
out of 18 old burrows examined, a litter was found in only one 
(Sdobnikov, 1958). 

In the arctic tundra of the Nizhne-Kolymsk region in the sum- 
mer of 1 950, the greater part of the burrows were inhabited, and 
in autumn, the number of arctic foxes was high. In 1951, the num- 
ber of arctic foxes shaфly decreased and out of 205 burrows, only 
11.5% of the burrows were inhabited; at the same time, in the 
southern zone of the tundra, where food is more varied, a concen- 
tration of arctic fox was noted. In summer of 1952, in the arctic 
tundra, out of 103 burrows 2.3% of them were inhabited, while in 
the southern 'Zone — 41.2%) (of 34 burrows). In summer and autumn 
of 1953, with abundant lemmings in the arctic tundra, more than 

Table 29. Occupation of burrows by arctic foxes in Turui^hansk 
territory, around 69-70° N. Lat. (Boitsov, 1937) 



Total 




Of these 


, in % 




burrows 
investigated 


With 3 
pups 


With signs 


Uninhabited 




Permanent 
residence 
of adults 


Rare 
visitation 




71 


2.7 


7.0 


11.1 


79.2 



368 




Щб % % '^^^ '^^' % % «^2 «/и <У.й <'/(« ^& %р Щ, Чь % % 

Fig. 91. Tanning of arctic fox skins in the Soviet Union as an index of 
fluctuations in the numbers of the species (A.F. Chirkova). 

40% of the burrows were occupied by arctic foxes and reproduc- 
tion proceeded well (average litters of 5— 7 young) (Tavrovskii, 1958). 
As is seen, the numbers of arctic fox do not change synchronously 
in the various geographic regions and even separate places, never- 
theless, great rises and falls in numbers simultaneously envelop vast 
areas of the European and Asiatic tundras eastward to the Lena 
[river]. Farther to the east, the rhythm of dynamics does not usually 
coincide. Only in occasional years the rise in number is observed 
in the area of the whole tundra zone of the Soviet Union, as it was 
in 1922/23, 1929/30, and apparently in the 3 years 1935/36-1937/38. 
A "crop" of arctic foxes in the winter of 1944/45 was noticed in 
the whole North with the exception of the Far East. 
256 . Even in the last century, the dependence of the numbers of 
mainland arctic fox on the abundance of lemmings, as the main 
food of arctic fox, was noted. At the present time, it has been 
convincingly shown that the massive appearance of lemmings stimu- 
lates reproduction in the arctic fox only when it takes place in the 
preceding autumn and winter (subnival reproduction of rodents). In 
the whole north of the Soviet Union, during the period 1944—1948, 
there can be traced a clear dependence of reproduction, migrations, 
and development of epizootics, and, as a result, the status of the 
numbers of arctic foxes, on the abundance of lemmings in the 
preceding autumn and winter, and of these rodents on the charac- 
teristics of weather in the course of different seasons of the year 
(Fig. 93). The value of these interrelationships and governing rules 
allows one to make a prognosis of numbers and population status 
of arctic foxes some months before the beginning of the harvest 



369 





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255 Fig. 92. Variations in relative numbers of the arctic fox in the tundras of 
the Soviet Union over 5 years — 1944/45-1948/49 (from Chirkova, 1955): 
1 — low number; 2 — number below average; 3 — average number; 4 — number 
above average; 5 — high [number]. 



370 




Moderate - 



Temperature 
Precipitation 



256 



Months I Ш 7Ш SS I'm 7'шаЛ~УШ'7ШПЛ1'ш'1ШКШ 1 Ш7 Ш SS 1ШТШ шл 

"""'^ ..J ШШШ1МЖ Ш71'Ш1Ж.ЕШШШ1Ж1 wn Mzini Ш и шит шш шил 
Years' тЗ J^ ' /54 J тб 



mi 



тз m9 



Fig. 93. Weather condition, reproduction of pestnishek* (lemmings) and 
number of arctic fox in the tundras of Yamalo-Nenetsk zone during the 

period 1943-1949 (from Chirkova, 1955). 

1 — numbers of arctic fox; 2 — ^numbers of lemmings; 3 — tanned arctic fox 

skins; 4 — prognosis of the number, based on sum of indexes; 5 — favorable 

weather (precipitation, temperature); 6 — moderate [weather]; 7 — unfavorable 

weather (for lemmings). 



season (Dubrovskii and Romanov, 1935; Romanov and Dubrovskii, 
1936; A. Romanov, 1941; Chirkova, 1951, 1955). 

Migrations have great significance in the dynamics of arctic fox 
numbers. As stated above, through this, emigration of huge num- 
bers of the animals from the tundra zone and their appearance in 
the taiga zone and on the Arctic Ocean, takes place. Latitudinal 
migrations of arctic foxes in some regions, for example, in Kanino- 
Timan tiandra at the time of great increase in their numbers, increases 
the total "crop" of arctic foxes, while the local groups of the 
animals remain small (Bashkin, 1940, Shibanov, 1951). On the 



* Local name — ^Sci. Ed. 



371 

Commander islands and the sea coasts of the mainland tundra 
257 in the east and west of the range in the USSR, where the food 
supply in the form of marine flotsam is relatively permanent, the 
number of arctic fox is more stable and changes less in the various 
years. 

Exploitation [for fur] also reflects numerical dynamics. Inten- 
sive trapping out of arctic foxes in years when they are abundant, 
increases the probability of survival among the remaining animals. 
Increased take of arctic foxes in the years when they are few in 
numbers is not rational, since it decreases the producer group. 
Hunting of arctic foxes in the burrows in the second half of winter, 
when they are preparing to breed, is particularly harmful. Rational 
utilization, taking into consideration the numbers of arctic foxes and 
the conditions of their existence, may favorably influence the popu- 
lation, increasing its productivity. 

Blue foxes, common in regions with mild climate, and extremely 
rare in mainland tundra (a tenth and a hundredth part of the per- 
centage among white arctic foxes; Paramonov, 1929; Lavrov, 1932; 
Zubkov, 1936 and others), are more often observed in years of 
abundance of the species. In Canada, blue arctic foxes also occur 
in larger quantities in years of massive reproduction of white arctic 
foxes (Hewitt, 1921). 

In Canada, massive purchases of skins are repeated each 3—5 
years, 4.2 years on the average. The catch of arctic foxes in 
individual years decreased 8-9 times (Seton-Thompson, 1909-1925; 
Hewitt, 1921). Co-occurrence of the year with an abundance of 
lemmings was established in both Norway and Canada (Elton, 1931, 
1931a). The numerical dynamics of the arctic fox in northern and 
regions of the Arctic and sub-Arctic zones of America differ. 
Increase in numbers of arctic foxes over large territories in Canada 
was noted in 1933/34, 1937/38, 1941/42 and 1945/46, and massive 
reproduction of lemmings— in the same or the preceding years (Elton 
andChitty, 1937, 1938, 1939, 1940, 1941, 1942, 1943, 1950). 

Stocks of arctic fox in Greenland are subjected to strong annual 
fluctuations depending on the arrival of spring, meteorological con- 
ditions which are determined by the amount and character of the 
passing of huge masses of drifting ice from the Arctic Ocean to the 
south along the shores of the island (Vibe, 1959). 



372 




Fig. 94. Track of an arctic fox on damp sand. Khatanga. Photograph by A.F. 

Chiri<ova. 

Field characteristics.^'^ The narrow paws of arctic fox leave 
tracks similar in form and dimensions to those of the red fox, but 
somewhat more blunt (Fig. 94). On the firm snov/ of the open 
tundra, the arctic fox moves about fi-eely, but in the soft snow of 
the forest zone it sinks in. Due to its highly fluffy paws, the con- 
tours of winter footprint are somewhat diffuse. The length of the 
track of the fore limb is 5-7 cm and of the hind, 5-8 cm. The track 
of female is smaller than that of the male. The load on 1 cm^ of 
the supporting surface of the limbs is 40-60 g. 

The tracks of all four paws are sometimes distributed sepa- 

258 rately or in pairs (Fig. 95). The usual movement of the arctic fox 

is a slow trot or easy gallop with the stride length from 45-50 cm 

to 100 cm or more. When pursued, the arctic fox develops greater 

speed. In times of fattening it, like the red fox, very thoroughly 

^^Middendorf, 1869; Nosilov, 1909; Cherskii, 1919-1920; Formozov, 1929, 
1946, 1959; Barabash-Nikiforov, 1937; Dubrovskii, 1937, 1940; Tyulin, 1938; 
L. Popov, 1939; Rutilevskii, 1939; Shastin, 1939; A. Romanov, 1941; N. 
Dement'ev, 1955; S. Popov, 1957; A.F. Chirkova. 



373 



щ 

pi 



Fig. 95. Tracks of a running arctic fox on a snow-covered surface. Khatanga. 
Photograph by A.F. Chirkova. 

searches all bushes, hummocks and projections above the soil sur- 
face. During the hunt, the arctic fox changes its gait, now slowly 
sneaks up, sniffing a track or following the prey at a distance, now 
creeping on its belly, then quickly glides forward or makes a sudden 
jump to overtake the victim. The arctic fox seeks out lemmings 
along their runways, senses and hears through the mass of snow. 



374 

vigorously digging them out even in the compressed snow up to 1 
m deep. At the capture site, only traces of blood or the stomach 
of the rodent remain. On Belyi Island in 1936, 49 holes 10-15 cm 
deep each were dug in the snow by an arctic fox in an area of 100 
m^ who made 3 passages in 50 cm reaching the soil itself; for all 
of this only one lemming was caught. 

In distinction from the red fox, the arctic fox makes broad 
trails, especially during mass movements along the seashore and 
259 river banks. On Novaya Zemlya, in years with abundant arctic 
foxes, one may, according to Nosilov (1909), travel by horse along 
their trails. On ice, the tracks of [polar] bear are often accompa- 
nied by the tracks of the arctic fox (Vrangel', 1841; Kiфichnikov, 
1937 and others). 

In the second half of winter, double and triple common trails 
and sometimes even tracks of four animals running together are 
often encountered. Traces of mating and fighting most often may 
be seen in the Lena-Khatanga territory in February; at this time, 
burrows emptied of snow are found. In the tracks of females, 
mucous secretions are met with — signs of estrus. After mating, 
arctic foxes lead a secretive way of life, and signs of their activity 
are rarely encountered at that time. With the bright sun of spring, 
the white arctic fox is difficult to distinguish on the snow. On the 
contrary, the blue arctic fox looks almost black. 

The breeding burrows are recognized by their well-worn* exits, 
the trampled vegetation, the digging and trails of the young arctic 
foxes, and the abundance of droppings and the remains of food. 
The presence of young in the burrow is betrayed also by the in- 
tense odor. Arctic foxes may be revealed by tapping on the soil 
surface above the burrow; young arctic foxes reveal their presence 
by short barks at such a time, while adults remain quiet. By the 
number of active entrances, one can judge the number of young. 

At the end of summer, after the dispersal of the litters, and in 
autumn at the time of migration of the arctic fox, the river banks 
and fast shore ice become dotted with their tracks. Before the 
rivers freeze or formation of fast shore ice in the sea, tracks of the 
animals are concentrated in a narrow 1-2 m strip along the edge of 
the water. With the growth of fast shore ice, the arctic foxes move 



*The Russian word obglazhennyc literally means either "picked," "gnawed," 
or "smoothed," "worn" — Sci. Ed. 



375 

with it farther away from the shore. They move out onto the 
thinnest ice, and only the sound of the ice cracking makes the arctic 
fox instantly jump backwards; during this, it raises its pads like a 
cat, fearing to wet them. To the side of the general path, only 
tracks of individual arctic foxes are observed, running hurriedly to 
a ravine or to the thicket of bushes, in order to search for food. 
From the tracks, one may see how the arctic fox can, with a 
sudden jump punch a hole in the thin ice to catch a small fish under 
it. A dog following an arctic fox onto newly-formed ice often falls 
through, while the arctic fox, being lighter in weight, only slips. 

The sense organs in the arctic fox are very well developed. 
The arctic fox can hear the movement of the small rodents under 
snow from some meters; it can react to bait at a distance of 300- 
400 m and run precisely in the required direction. It can see willow 
ptarmigan far away, entirely invisible on the white snow cover of 
the flat tundra. The sense of smell is excellently developed in the 
arctic fox; it can scent a capercailzie in the snow at 100 m. When 
the animal seeks food, its head hangs down and the nose "traces" 
the path. In its absoфtion in the search, especially when young, the 
arctic fox often does not pay any attention to surrounding objects 
and approaches a stationary human to within 2—3 m. 

The arctic fox does not belong in the number of silent animals, 
and gives voice in various situations. The sounds given by it are 
quite varied. The usual cry of the arctic fox resembles that of a 
dog's bark, but more hoarse, deep and husky. An arctic fox hidden 
among rocks grumbles "urin-urin". Caught during migration, sitting 
in a sack, the arctic fox gives a cry "ugkhu-ugkhu" when ap- 
proached by a human. During the time of rut, females howl, yelp 
and squeal. When young arctic foxes disperse from the burrow 
prematurely because of lack of food, the agitated parents call them 
with a cry similar to that of the swan; this is most often heard at 
dawn. Hunters say "The arctic fox coos"; such a cry is consid- 
ered to be a forerunner of migration. 

On Novaya Zemlya, in the autumn of 1887 and 1888, the bark 
of arctic foxes was often heard at the time of migration. While 
running it might pause, and, raising its snout, it barked not strongly, 
but "complainingly". Arctic foxes also howl while running in win- 
ter on lake ice. Judging from everything, howling in arctic foxes is 
related to deficiency of food. (A. Ch.). 



376 

260 Practical Significance 

The arctic fox is one of the valuable species of fur-bearing animals. 
The USSR occupies first place in the world in the size of its fox 
fur trade. In pre-revolutionary times, tens of thousands were ob- 
tained in Russia and about 3/4 of the harvested catch was sold in 
internal fairs, of which the chief were the Irbitsk, Nizhegorod and 
Yakutsk. Part of the skins were exported to China, Manchuria, 
Japan and America. There is information on the exportation of 
great quantities of arctic fox skins from some regions of Russia. In 
the middle of the 18th century, according to Steller, more than 7000 
arctic foxes were caught during three wintering periods on Bering 
Island. In 1825 and 1826, 140 thousand skins were exported from 
Turukhansk territory; from Mangazeya, up to 40 thousand. Indi- 
vidual authors refer to this in connection with the decline in the 
harvest of arctic fox. The decline in catch of arctic fox was par- 
ticularly shaф in the last years of the Tsarist government. 

Organized exploitation of arctic fox in the USSR began in 1922 
(Paramonov, 1929; Guber, 1939; A.F. Chirkova). During the last 35 
years, tanned arctic fox skins in the USSR consisted of an average 
of 78 thousand per year, ranging from 35 to 128 thousand. From 
1946/47 to 1958/59, the average amount of skins tanned in a year 
increased to 8 1 thousand. This was promoted by the utilization of 
new areas, provision of supplemental feeding* of animals, the tak- 
ing and holding in captivity of migrating arctic foxes, and also the 
rationalization of methods and tools of exploitation. 

As regards the price of furs, before the October Revolution, 
the arctic fox occupied second to fourth place after squirrel, sable 
and red fox, depending on the "harvest" (Nosilov, 1909; Guber, 
1929; Chirkova et al., 1959). For the last ten years, the arctic fox 
stands in second to third place, providing 15% of the price of fiirs 
tanned in the USSR, while in the tundra zones — up to 90-99%, and 
representing the main object of hunting trade for native peoples of 
the Far North. The most valuable white fox ftir is obtained from the 
Yenisei tundras, Yakutiya and Novaya Zemlya. The main regions 
of exploitation are the Ob', Yenisei and northern Yakutiya; during 

261 the last 20-30 years they provided 30, 25 and 23% respectively of 



*See footnote. Table 25 — Sci. Ed. 



377 

all of arctic fox production in the USSR. Arctic foxes caught in 
northern Arkhangel'sk district (14%) and in the Far East (7%) are fewer. 

During the above-mentioned 30-35 years, an average of 5.8 
arctic foxes are caught annually in an area of 100 km^ of the 
tundras and forest-tundras in the Nenetsk and Yamalo-Nenetsk 
national regions, in Krasnoyarsk territory — 3.2; in Yakutiya — 3.7, 
and in the Far East — 1.2. At the same time, in the territory of the 
kolkhoz [collective] "Krasnyi Oktyabr' " [Red October], on the 
Kara Sea coast, in the 1950's, an average of 19 arctic foxes were 
caught yearly, and in years with a good "harvest" — ^up to 30. In the 
end of the past and at the beginning of the current century, on the 
Commander islands, up to 13 arctic foxes per 10 km^ were cap- 
tured by means of drives (through the year). Success of the harvest 
of arctic foxes differs greatly by year. One hunter taking 10-20 
arctic foxes in a season of moderate numbers, may catch up to 100 
or more animals in years of abundance. Individual hunters in north- 
eastern Yakutiya obtained up to 176 arctic foxes, on Wrangel' 
Island — up to 125, and in northeastern Bol'shezemel'sk tundra — 
more than 200 arctic foxes per season. 

In connection with the shaф variation in the numbers of arctic 
foxes, there arose the necessity for an organized service to esti- 
mate the "harvest" and to make a prognosis of arctic fox number. 
It was organized by the Administration of Glavsevmoфut' [Main 
Northern Sea Route] in 1935, when the first general prognosis of 
changes for the whole North was given, which proved to be true 
for a considerable part of the USSR (Dubrovskii and Romanov, 
1935; Romanov and Dubrovskii, 1936; Guber, 1939). The institute 
of the hunting industry (VNIO) [All-Union Scientific Research 
Institute of Hunting], in the course of a series of years, presented 
an analysis of the condition of the populations for the tanning 
organizations, and gave a prognosis of the number of arctic foxes 
in all sectors of our Far North; they were verified in the over- 
whelming majority of cases (Chirkova, 1955). 

At the present time, the main methods of arctic fox capture 
combine old ways and equipment with new techniques. In some 
places, the old method of capture are used; driving with [rein]deer, 
barriers on the banks of rivers and the sea, but they are additionally 
equipped with new types of deadfalls, food-baits, the newest jaw- 
traps and explosive weapons. Wooden deadfall traps are mainly 
used in the Taimyr National Region and in Yakutiya, and rarely in 



378 





SJefci^W^isafs^^iSfc;-' - 






260 Fig. 96. Slopets, or deadfall, for arctic fox (side wall open) in northeastern 
Bol'shezemel'sk tundra (European North). Photograph by Yu.V. Rybalkin. 
[See also Fig. 97.] 

Other tundras (Figs. 96, 97). They represent a narrow corridor with 
a bait in a box.* The arctic fox is killed by a log, hung above the 
corridor. Deadfalls are located along the seashore, river banks and 
in other sites, where the arctic foxes are most frequently encoun- 
tered. On the Taimyr peninsula, trap-lines extend from the border 
of the forest in the south northward deep into the tundra for 100- 
262 150 km and more. At the present time, improved traps are being 
deployed, especially portable traps — ^irreplaceable equipment for har- 
vest in sparsely populated tundras. The steel jaw-traps occupy 
second place among the implements of trapping. They serve as the 
main trap used for harvesting arctic fox in the western tundras and 
in northeastern Siberia. 



''The Russian word is strozhka, literally "lodge" or "hut" — Sci. Ed. 



379 







!;■■ 




Fig. 97. View of deadfall from the front. Photograph by Yu.V. Rybalkin. 
[See also Fig. 96.] 

A pot drive*, or "ro/ar" is employed on Yamal and in the 
tundras of the European North. In it, from 2-6 tens of hunters on 
reindeer teams participate, travehng in a spiral over open, compara- 
tively level territory of an area of some tens of square kilometers. 
Arctic foxes, fearing to cross the fresh trail of the deersled, gather 
in the center of the site around which the harvesters are traveling. 
The hunters kill the arctic foxes with guns, or even sticks. In years 
of abundance they take up to 1 50-200 arctic foxes in a circuit. The 
method of tolars on foot also exists. Sometimes, the arctic foxes 
are driven on to promontories of the seashore or into nets that are 
setup (Romanov-Il'inskii, 1958; Skrobov, 1963). 



''The Russian phrase is zagon kotlom — Sci. Ed. 



380 




Fig. 98. Blue arctic foxes arrived for supplemental feeding. Mednyi Island. 
December 1953. Photograph by S.V. Marakov. 

Arctic foxes are frequently shot from a blind, during solitary 
stalks on [reinjdeer or horses, with dogs or in an accidental encoun- 
ter. Harvest of the arctic fox on the seashore during the time of 
autumn migration is permitted only in an organized arrangement for 
the sake of keeping the animals in captivity until full pelage. For- 
bidden everywhere is klyapts*, digging out animals from burrows, 
baited crossbow-traps, and baits poisoned with strychnine, corro- 
sive sublimate [mercuric bichloride], nuxvomica or poisonnut, 
tormentil and other poisons. The modem method of catching live 
arctic foxes by means of food-supplied live traps is employed mainly 
in the island enteфrises, but has begun to be used in mainland 
tundras. In Bol'shezemel'sk tundra, portable food-supplied live traps 



"Meaning not clear; perhaps a kind of trap — Sci. Ed. 



381 

of simple construction are used (Boitsov and Pereleshin, 1934; 
Freiberg, 1939; Skrobov and Kozhevin, 1955, 1957). 
263 The best methods for catching the arctic fox are snares and 
traps based on the employment of lures and baits. Lures for the 
arctic fox may be visual, food, scent and mixed. Each object on the 
snowy surface of the tundra can be seen from afar and attracts the 
arctic fox's attention, and therefore hunters drive into the snow 
pieces of deer-antlers, chocks, set up piles of stones, and pile up 
hillocks of earth or snow, etc. Food lures are very varied. However, 
fresh carcasses of lemmings, grouse, hares and spoiled eggs are a 
continual preference of the tundra population of arctic foxes. Arctic 
foxes of the coastal regions prefer fresh fish and fat of marine 
animals. Blood of freshly-killed [rein]deer and other animals acts as 
a good lure. In order to attract arctic foxes to hunting areas before 
the beginning of the harvest, they place baits such as carcasses of 
common seals, ringed seals, white whales and other animal and 
fishing industry wastes to which arctic foxes are attracted to feed. 
This considerably increases the catch. 

In previous times, the hunt for arctic foxes continued through 
almost the whole year. In the middle of summer the young nornik 
were dug out of burrows for nursing. The regular harvest of skins 
began at the end of summer, taking in abundance krestovatik 
[cross], sinyak [blue] and nedopesok [premature], and the catch 
continued to April. In years with an abundance of migrating arctic 
foxes, a coastal harvest existed, when 5 to 10 or more animals not 
in full pelage were taken per night. In such years, more than 50% 
of the harvest consisted of "nevykhodnye"* arctic foxes. In the 
past, arctic fox skins were given as tribute, from each Nenets two 
arctic foxes were taken, while from Evenk and Dolganin — four. As 
a quantitative unit of exchange, among the Vaddeevsk Samoyed, 20 
arctic foxes were equal to one boat. 

From the first years of the Soviet state, the catch of 
nevykhodnye arctic foxQS— nornik [burrow dwellers], krestovatik 
[cross], sinyak [blue] and the spring ''gagara" [loon] was forbid- 
den. A harvest season was established from the middle of 
November-beginning of December, to the beginning or end of March 
(in various regions). 



^Literally "unfinished", i.e. immature — Sci. Ed. 



382 

An arctic fox industry is conducted on islands surrounded by 
unfrozen seas (Boitsov, 1937; Il'ina, 1949-1950 and others). Bering 
Island supplied arctic fox to Andreyanov Island (Aleutians) which 
belonged to Russia at that time (Khvostov and Davydov, 1810- 
1812). At the end of the eighteenth century, G.L. Pribylov [Pribilof] 
exported 8,000 skins of blue arctic foxes from Georgii and Pavel 
[St. George, St. Paul] islands in two years, and only after that arose 
the question of the preservation of these animals there. It was of 
great practical importance to organize an intensive arctic fox indus- 
try on the Commander islands, where blue arctic foxes occur. In 
1741, when Bering visited these islands, blue arctic foxes thrived 
there in "incalculable packs". Steller killed tens of arctic fox with 
an axe in a day, and used the skins of the animals to cover the 
roofs of the earthen huts. At the end of the 90's of the nineteenth 
century, 1.5 to 2.5 thousand arctic fox skins were taken annually 
from the Commander islands. In the beginning of the twentieth 
century, as a resuh of overharvest, drives began to be allowed only 
in alternate years, preserves were allotted, counts of arctic foxes 
were promoted to determine the number allowed to be killed, and 
the permissible period of capture was shortened to 2—3 weeks. As 
a result of many years of selection, white arctic foxes, individuals 
of which appeared on Bering Island particularly often, were exter- 
minated. Only under the Soviet government in the 20's was winter 
supplemental feeding introduced and in the 30's, in summer as well. 
Food-baited live traps began to be used and continuous zootechnical 
selection conducted. It is supposed that the arctic fox population on 
the Commander islands can be increased to 40 thousand head. In 
the 20' s, attempts were made to transfer arctic foxes to the small 
coastal islands of the Pacific Ocean and to the Solovets (united in 
winter with the mainland by an ice bridge), with the aim of orga- 
nizing there an island blue arctic fox industry. All of these ended 
unsuccessfully. 

Experiments on supplemental feeding of the arctic fox showed 
the great importance of this measure not only for islands, but also 
in the mainland tundra on several farms in isolated territories. Supple- 
mental feeding must be begun during the time the litters are still in 
264 the burrows — ^at the first signs of the disappearance of natural food 
in spring, summer and early autumn, to inhibit the arctic foxes from 
migrating, and also in winter in places where animals concentrate 
and before the beginning of rut. 



383 




^^,^^' 



Fig. 99. Blue arctic fox lying in rocks. Bering Island. October 1958. 
Photograph by S.V. Marakov. 

Intensive arctic fox farming on islands has been conducted in 
North America for the past century, based mainly on wastes of the 
marine animal industry (Suvorov, 1927; Boitsov and Pereleshin, 
1934; Boitsov, 1937). As already mentioned, the first release of 
blue arctic foxes on the Aleutian chain was done by Russians in the 
middle of the eighteenth century already, and on the coastal islands 
of Alaska in 1835, and American trading voyages for this activity 
were undertaken only in 1885. Supplemental feeding of arctic foxes 
on the Pribilof islands began in 1896; later other measures were 
carried out. Concentrations of arctic foxes here attained very high 
densities; up to 500 arctic foxes occurred for each food-baited live 
trap. 



384 

Commercial raising of arctic foxes in state animal sovkhoz* 
was begun in the USSR in the 30' s. Raising the blue form was 
introduced rationally due to their great value. Cage raising of blue 
arctic foxes, developed in a series of European countries and in 
North America, past through a crisis in the post-war years as a 
result of the creation of a mode for short fur. At the end of the 
1950's, some rise in demand was again noted, though public taste 
turned to white and light-blue arctic foxes. 

The proportion of white arctic fox in the fur export of USSR 
reached 7.3% (1933). Before the Great Fatherland War**, blue 
arctic fox was priced significantly more expensively than white in 
the world market, and the darker, the higher, but in the Leningrad 
fur auction (1961), white arctic foxes sold with higher prices than 
the blue, and among the latter, the more expensive skins were of 
light color. 
265 World production of arctic fox fur (Brase, 1925) in the period 
1907—1909 averaged 1 16.0 thousand skins, of which 105.0 thousand 
were white, and in 1923—1924, 204.0 thousand skins, of which 166.0 
thousand were white. North America contributes on average about 
1/3 of the skins of white arctic fox. In the 20's and 30's, from 18 
to 72 thousand skins were bought in Canada, an average of 44.9 
thousand. In Greenland at the end of the last century, an average 
of 1.5 to 2.0 thousand arctic foxes were caught every year. In 
Spitsbergen in the 19th century, the arctic fox was exploited espe- 
cially intensively. In the 20' s of the current century, only 100 blue 
and 250 white arctic foxes were obtained annually. At the present 
time, great attention is paid there to the preservation of arctic fox 
(Soyrinki, 1958; Bergman, 1961). 

Arctic fox meat has an unpleasant odor, but on Yamal many 
people ate it (Nosilov, 1909; N.N. Spitsyn). At the present time, the 
question of the specific activity of the fat of arctic and red foxes 
as a medical dietary treatment against tuberculosis and liver dis- 
eases in humans arises (Mironov, 1953). 

Arctic fox has a negative significance as a disperser and trans- 
mitter of tundra rabies. In years of outbreaks of this disease, many 
sled- and herd dogs and reindeer are bitten by arctic foxes, are 
infected and die. However, the frequency of disease transmission 



* State farms — Sci. Ed. 
**World War II— Sci. Ed. 



385 

by arctic foxes may be exaggerated due to insufficient study of the 
significance of rodents and other animals of the tundra. Since 
echinococcosis, trichinosis, diphillobothrosis, opistorchosis and other 
helminthoses are found in arctic foxes, the possibihty of the trans- 
mission of these diseases to humans is not excluded. (A. Ch.). 

Genus of [Common] Foxes 

Genus Vulpes Oken, 1816 

1775 Vulpes. Frisch. Natur-Syst. d. Vierfiiss. Thiere, p. 15. By 
decision of the International Committee on Nomenclatural, 
this work was considered to have no nomenclatural signifi- 
cance.* 
1816 Vulpes. Oken.* Lehrb. d. Naturgesch., 3, Th. 2, p. 1033, 

1034. Vulpes communis Oken = Canis vulpes Linnaeus. 
1839 Cynalopex. H. Smith. Jardine's Nat. Library. Mamm., 25, 
p. 222. Canis corsac Linnaeus (V.H.). 
Species of medium and small size. 

Skull light and slender, elongated, with widely situated zygo- 
matic arches, and low (height in occipital region less than 1/3 of 
condylobasal length). Skull sculpturing is weakly defined, sagittal 
crest not developed at all or weakly defined only in posterior part 
of the parietal bones. Facial part light and slender, considerably, 
sometimes very strongly elongated and much longer than braincase; 
its length from the posterior wall of infraorbital foramen to posterior 
edge of the canine alveoli greater than skull width above the ca- 
nines, while distance from line connecting ends of supraorbital 
processes to anterior end of premaxillae significantly greater than 
that to upper edge of occipital foramen. 

Interorbital (frontal) region flat, not swollen between supraor- 
bital processes — ^here a concavity (groove) is located; supraorbital 
processes large, dorsally somewhat concave, with small air cavities, 
a quite shaф edge and with apex directed somewhat downwards. 
Profile line of rostral part of skull grades very gradually into frontal- 
parietal line, not forming a noticeable, or only an ill-defined, projection; 
sometimes, upper profile of skull conforms almost to a completely 
straight line. 
266 Posterior edge of tooth row reaches level of anterior edge of 
interpterygoidal fossa. No deep notch and projection on lower edge 

*However, the International Commission on Zoological Nomenclature (1979) 
retained Vulpes Frisch 1775, while rejecting Vulpes Oken 1816. 



386 

of lower jaw in front of angular process. 

3 14 2 
Dental formula I— C-P— M— = 42. In some, as individual de- 
3 14 3 

3 14 2 
viation it may be I— C-P— M— = 40. Teeth relatively small and 

^3142 ^ 

light, with very shaф cusps and ridges. Length of upper camassial 
tooth and molars 2%— 3 times more than palatal length. Canines thin, 
strongly curved and very long — ^when jaws are closed, tips of upper 
canines reach or nearly reach lower edge of mandibular bone or 
even extend beyond it, while tips of lower canines are located 
above edge of alveoli of upper canines (Fig. 18). Ends of upper 
canines directed straight downwards; no cutting edge develops on 
posterior side of canine; canine surface entirely smooth. Incisors 
usually without accessory lobes. 

Trunk long, limbs quite short, tail very long — more than half that 
of body, always falls below tarsal joint, and in standing animal 
reaches the ground. Head elongated, with long pointed muzzle. 
Ears long, when bent forward always reach eye, sometimes very 
long with pointed tip. No large bare callosity found on the rear 
extremities behind four bare toe pads (a small one occurs — feet 
covered with fur). Glandular area is found in front of the middle 
callosity on the fore and hind limbs. 

In some, fiir dense, fluffy, long and soft (northern forms), in 
others, more sparse and coarse. Color monophasic, in one species 
(red fox, V. vulpes) polyphasic, and, except for black phase of this 
species, adults do not exhibit only one color. In various species, the 
prevailing colors are reddish, sandy- and clayey-yellow and brown 
tones with silvery-white and black in the form of admixtures and 
black markings. Supracaudal gland well developed. Nipples 4 or 6 
pairs. No marked sexual dimoфhism (males are slightly larger), age 
dimoфhism well manifested and seasonal dimorphism is shaф in 
some forms (northern), but is revealed almost solely in degree of 
fur density and length. There are two molts per year. 

In general appearance, species of the genus are relatively very 
топотофЬ1с, differing mainly in size. In this respect, three types 
occur: the relatively large red foxes {V. vulpes), forms of lesser 
size — corsac fox {V. corsac) and its analogues, and very smaU 
foxes of the Baluchistan type {V. сапа). The largest forms have a 



387 

body length up to 90 cm and weight up to 14 kg and the smallest — 
about 40 cm in body length and hardly more than 2 kg in weight. 

Tliey are inhabitants of extremely varied landscapes — from tun- 
dras and taiga to extremely high mountains, and steppes and deserts. 
They prefer areas with more or less open expanses and avoid the 
dense thick forests. There are no regular migrations. They are 
monogamous, uniting for one reproductive season. Up to 13 young, 
usually less, bom in burrows. They are carnivorous animals, feeding 
on small vertebrates, mainly rodents, carrion, and as an exception, 
on insect and plant food. Except for the breeding season, they lead 
a solitary way of life. 

The range of the genus is very vast, located on four continents 
and in all climatic zones. It occupies all Europe, to North Cape and 
islands of the Mediterranean Sea, includes the British islands (ab- 
sent in Iceland), all Africa except Madagascar, and Asia northward 
to the northern shores of the continent (except the northern Taimyr) 
and the Kolguya islands, Yuzhnye Novaya Zemlya and Belyi is- 
lands (locally wandering animals may be seen farther north; see 
268 description of red fox, V. vulpes) and southward to Tonkin, Yunnan, 
Sichuan, Assam, Bengal and India to Cape Comorin. It occupies 
the entire Arabian Peninsula. The range includes Karagin, St. 
Lawrence, Kuril, Sakhalin, Shantar and Japanese islands. Species 
of this genus are absent on the islands of the Bering Sea (except 
those mentioned). 

In North America, the range extends northward to the northern 
coast of Labrador, Hudson Bay, a line from Chesterton (on the 
western shore of Hudson Bay) to Coppermine (on the Arctic coast), 
and farther westward on the Arctic Ocean and Bering Sea coasts. 
Southward, the range extends to California and northern Mexico 
inclusive, but does not include the southern and southeastern states 
of the USA from Maine to Louisiana (for details, see range de- 
scription of red fox, V. vulpes). The range includes Newfoundland 
and Kodiak islands. Species of the genus are absent from the 
coastal belt of British Columbia and adjacent islands. Acclimatized 
in Australia {V. vulpes). In the vertical direction, they reach ex- 
treme heights. The range of the genus was not subjected to 
substantial reduction in the past century. 

In systematic characteristics, Vulpes represents a more spe- 
cialized group than Canis and Alopex, and the degree of specialization 
of some species (V. ferrilata, V. сапа) is very high. Besides the 



389 

close relationship to the arctic fox, Alopex (see above), Vulpes is 
closely related to the fennec group — genus Fennecus. All three 
genera are considered "neighboring" groups. Through Alopex, 
Vulpes is related to Canis (wolves and jackals) (see characteristics 
of family and of named genera). The relation to genus Ducicyon 
(American gray fox and close species), with which the described 
genus is sometimes associated is, apparently, more distant. Since 
the extreme forms of all the genera mentioned have much in com- 
mon, and serve as connecting links between them, there is basis for 
considering all of them as only subgenera of one broadened genus — 
Canis (see the family characteristics). At the present time, however, 
it is still almost always considered as qualifying as independent. 

The genus already was separate in the Pliocene and may have 
appeared even in the lower Miocene (North America). In Europe 
and Asia, it is known in the Pleistocene. The genus Vulpes, like 
Canis, probably developed from the American Mio-Pliocene group 
Tomarctos. 

In the contents accepted here, the genus is quite monotypic and 
was not subjected to any considerable subdivision. Attempts were 
made to split into a separate genus the corsac fox {V. corsac) and 
unite it (sometimes also with V. velox) with genus Alopex (arctic 
foxes). 

In the genus are 6 species: 1) red fox, V. vulpes; 2) corsac fox, 
V. corsac; 3) Bengal fox, V. bengalensis; 4) Afghan fox, V. сапа; 
5) American small fox, or American corsac, V. velox* and 6) 
Tibetan fox or Tibetan corsac, V. ferrilata. The genus includes 
aboui 20% of the species of the family, 4 Asiatic species (F. 
corsac, V. bengalensis, V. сапа, V. ferrilata), 1 American spe- 
cies (F. veloxY and 1 species distributed in the greater part of 
North America, in almost all Eurasia and in North Africa {V. vulpes). 

The African V. chama and the African-Near Eastern V. 
rueppelli are sometimes included in genus Vulpes; however, they 
are apparently to be referred to genus Fennecus. 

The fauna of the USSR includes 3 species: 1) red fox, 
V. vulpes Linnaeus, 1758; 2) corsac fox, V. corsac Linnaeus, 
1768; 3) Afghan fox, V. сапа Blanford, 1877, i.e. 50% of the 



*In English, swift fox; V. megalotis, the kit fox, is often considered a distinct 
species as well — Sci. Ed. 



390 

species of the genus, and about 1% of the species of the USSR 
fauna'. 

The range of the genus covers the entire territory of the USSR, 
except the above-mentioned northern regions. Species of the genus 
are met with in all landscapes of the country. 
269 The members of this species are useful as commercial fur- 
bearing animals, and in part for limiting the number of harmful 
rodents. In places, they cause some harm by killing useful sporting 
animals and birds (V.H.). 

AFGHAN FOX^ 

Vulpes сапа Blanford, 1877 

1877. Vulpes сапа.* Blanford. Journ. As. Soc. Bengal., 46, pt. 2, 
p. 231. Gwadar, Baluchistan (Mekran coast). 

1907. Vulpes сапа var. nigricans. Shitkow. Zool. Anz., 32, p. 
448. Southern Tadzhikistan ("Gomaya [montane] Bukhara"). 
V.H. 

Diagnosis 

Dimensions very small, body length not more than 50 cm, 
condylobasal length of skull not more than 91 mm. Tail tip not 
white, back color gray mixed with black ripples, without reddish and 
brown tones, or they are very weakly developed (V.H.). 

Description 

The Afghan fox is a carnivore of typical "foxy" appearance, but 
of very small size. It has relatively short legs, elongated trunk and 
very long tail. Its length is considerably more than half the length 
of the body, and in a standing animal it reaches the ground. Head 
with very pointed, typically fox-like muzzle and large, wide, erect 
pointed ears. Their length more than half the length of hind foot. 
Number of teats unknown. 



Mnformation on the occurrence of V. nieppelli Schinz, 1825 ["F. (Megalotis) 
famelicus Cr."; Flerov, 1932] in southern Turkmeniya is a mistake. 
^''Shardel'" of furriers. 
* Misspelled "canus'" in Russian original — Sci. Ed. 



391 




Fig. 101. Afghan fox, Vulpes сапа Blanf. Sketch by A.N. Komarov. 

In winter, the fur is dense, fluffy, very soft and relatively quite 
long. Dorsal guard hairs 45-65 mm long, underfur, 35-45 mm. Tail 
covered with dense, long hairs (their length 65—90 mm) and very 
fluffy. 

General tone of winter fur brownish gray, with well-developed 
black highlights along back originating from guard hair color. Guard 
hairs have black terminal portion, with a small silvery-white zone 
proximal it. Some g\iard hairs are entirely black. Guard hairs form 
on the skin a silvery frosting. On head, general color lighter and 
somewhat whitish. Well-defined blackish-brown stripe runs from 
comer of eyes to base of vibrissae. Light rusty frosting develops on 
occiput. Rear sides of ears gray in color or blackish-gray. Chin 
270 dark brown or brownish-black. On sides of the body, color consid- 
erably lighter than on dorsum, changing into pale yellow-gray. Here 
admixture of black guard hair is small; guard hairs have gray and 
silvery tips and underfur lighter than on back. Throat, lower part of 
neck, chest, and whole abdominal surface white, with light straw- 
colored tones. Extremities gray in color, more or less dark, with 
light rusty highlights on external side. On fore feet, black markings 



392 



sometimes present. Tail gray, with strong blackish highlights above. 
In region of supracaudal gland, on upper side near base, is a well- 
defined dark spot. Underftir is mouse-gray in color, their tips with 
light, or more or less intense ocherous highlights. 

Individual variation in the color described above are known. It 
is expressed chiefly in that individual specimens are tinged espe- 
cially dark gray in color without any ocherous or rusty admixture 
or with a very weak admixture of these tones, above all on the legs. 
In others, the admixture of ocherous and rusty ("foxy") tones is 
more noticeable, and the rusty is well expressed on the shoulders. 

Summer pelage is not described. The first [juvenile?] pelage 
and age variation in color are not known. 

The fox we are describing is very rare in collections and its 
amplitude of variation is poorly known. Generally, this species is, in 
many respects, poorly studied. 

Skull resembles that of corsac fox, but facial portion relatively 
shorter, braincase relatively larger and more expanded. Zygomatic 
arches widely separated and are quite broad anteriorly, from which 
its general outline is more or less rounded (together with rostral 
portion, not forming evenly tapered clinoidal figure). Nasal bones 
narrow and highly tapered towards posterior end. Sagittal crest not 
developed, and temporal lines stand far apart from each other, 
forming a lyriform figure. Molars relatively large; canines thin and 
long. Middle incisors without accessory blades. 




Fig. 102. Skull of Afghan fox, Гм/ре^сало Blanf. (after Pocock, 1 94 1 ). 



393 

Chromosome number unknown. 

Size very small — this is the smallest species of the genus in our 
fauna, and in the genus generally. Body length 40-50 cm, tail length 
33-41 cm, hind foot about 100 mm, ear length about 90 mm. 

Condylobasal length of skull, 89-91 mm; zygomatic width, 50- 
51 mm; interorbital width, 18—19 mm; length of hard palate, 45 mm; 
length of nasal bones about 29 mm, length of upper molar row, 41— 
44 mm; length of lower jaw, 67—69 mm; length of upper canine 
about 10 mm; and of lower, 8 mm (according to Pocock, 1941, and 
Novikov, 1956; published data are those of separate individuals; in 
our museums, there are no fully valuable materials). 

Information on comparative sexual dimoфhism in size is absent; 
as in all foxes, females are probably smaller than males (V.H.). 

Geographic Distribution 

Eastern part of Near East. 

Geographic Range in the Soviet Union 

This represents the extreme northern periphery of the species range. 
In dimensions it is insignificant and is limited to the extreme south 
of Middle Asia. 
271 Reliable places of capture of the Afghan fox lie in the Kopet- 
Dag at the northern foot of the western part of the Sarymsakli 
range, 40 km from Germab, which is west of Ashkhabad (1926; 
Ognev, 1931), and at Bakharden on the railway line between 
Ashkhabad and Kyzyl- Arvat (1926; Novikov, 1956). 

In the 90's*, 4 skins were brought to Moscow from 
"Vostochnaya [East] Bukhara", i.e., southern Tadzhikistan (Zhitkov, 
1907); one of these specimens is preserved in the Zoological Mu- 
seum of Moscow University in stuffed form. At that time, the 
possibility was admitted that these were not native, but imported 
from Afghanistan. The majority of later authors shared this belief. 
At the present time, information has appeared that skins of this 
species occasionally come in from southern Turkmeniya and 
Tadzhikistan (Kuznetsov, 1952). According to other data 
(Chemyshev, 1958), "during the course of a prolonged period", not 

*In Russian original, "V 900-kh godakh" [In the 900's; i.e. 10th century]. 
This evidently refers to the 1890's — Sci. Ed. 



394 

once has a skin of this species appeared among the fur prepared 
in Tadzhikistan, Thus, the question on the occurrence of Afghan 
foxes on the eastern shore of the Amu-Dar'ya and Pyandzh re- 
mains open and, most likely, will be settled in the negative. 
Southwestern Turkmeniya remains the established area of its oc- 
currence within the borders of our country. 

Apparently, one must now reach the conclusion that the 
Afghan fox in our country is only vagrant, appearing rarely in the 
Kopet-Dag and adjacent plains. This corresponds to the known 
occurrences of the species in northeastern Iran (Khorassan; Duruh 
in the eastern part of the province). It may be assumed that the 
species under description appears also in southeastern Turkmeniya — 
along the Tedzhen and between Tedzhen, Murgab and Amu-Dar'ya 
[rivers]. These areas lie south of the places of transience of the 
Afghan fox to the west [Fig. 103]. Wandering into Tadzhikistan is 
also doubtful because the Pyandzh and Amu-Dar'ya are probably 
insuperable barriers for such a small animal. 

Geographic Range outside the Soviet Union 

This includes Afghanistan, the northwestern provinces of India 
bordering Afghanistan, Sind, Baluchistan and eastern Iran, in par- 
ticular Khorassan. 
272 The range of the Afghan fox is notable for its very small size, 
which is unusual for a carnivore, and particularly for the species of 
the family Canidae. It is even smaller than the range of the rela- 
tively narrowly distributed Bengal (V. bengalensis) and Tibetan (F. 
ferrilata) foxes (V.H.).* 

Geographic Variation 

Geographic variation in this species is not described, and probably, 
is not manifested — its range is too small for this. Moreover, it is 
restricted to one natural geographic landscape region. Based on all 
data, the above-mentioned specimen described from "Vostochnaya 
Bukhara" as "var. nigricans'' represents only an individual color 
deviation (V.H.). 

* V. сапа has recently been found in northwestern Iran, the Arabian peninsula 
and Israel (Harrison and Bates, 1991), where it occupies steep, rocky habitats — 
Sci. Ed. 



395 




271 Fig. 103. Range of the Afghan fox, Vulpes сапа Blanf, in the USSR. Dots 
designate places where animals were obtained, question marks — areas 
of assumed occurrence. V.G. Heptner. 

Biology 

There is no information whatsoever about the biology of the 
Afghan fox within the USSR. Data from other countries where this 
species Hves, especially India and Pakistan, are also absent (V.H.). 



273 



Practical Significance 

In our country, the Afghan fox has no practical significance at all. 
As for the small number of skins, "shardeF ", which sometimes 
appear among lots of furs from Tadzhikistan and Turkmeniya, and 
probably from the neighboring countries (see above) they do not 
play any role whatsoever in fur trade. Apparently, the agricultural 
significance of this species is also negligible. No substantive mate- 
rials are available from the other countries (V.H.). 



396 




272 Fig. 104. Species range of the Afghan fox, Vulpes сапа Blanf Very schematic. 

V.G. Heptner. 



CORSAC FOX 



Vulpes corsac Linnaeus, 1768 



1768. Vulpes corsac. Linnaeus. Syst. Nat., Ed. XII, 3, App. p. 
223. Steppes between Ural and Irtysh rivers. Restricted to 
"Pri-Irtysh steppes south of Petropavlovsk" (Ognev, 1935). 



397 

1912. ? Vulpes corsac var. nigra. Kastschenko. Kashchenko. 
Ezhegodnik Zool. muzeu I. [Irkutsk] Akad. Nauk, 17, p. 393 
Trans-Baikaliya. 

1935. (1934)-* Vulpes corsac scorodumovi. Dorogostaiski. 
Dorogostaiskii. Izv. Irkutsk, gos. protivochumn. inst. Sibirii i , 
D.V. kraya. 1, p. 47. Southern Trans-Baikaliya. 

1935. Vulpes corsac kalmykorum. Ognev. Zveri SSSR i prilezh. 
stran [Animals of the USSR and adjacent countries]. 3, p. 
634. Kalmytsk steppes (steppes on right bank of Lower 
Volga; V.H.). 

1935. Vulpes corsac turcmenicus . Ognev. Zveri SSSR i prilezh. 
stran [see above] 3, p. 635 "Turkmen and Tadzhikistan 
semideserts and deserts". It is proposed to restrict this to: 
Murgab and Tedzhen districts and lowlands at foot of east- 
em Kopet-Dag (east of Ashkhabad). V.H. 

Diagnosis 

Of moderate size, larger than Afghan, but smaller than red foxes. 
Body length more than 50 cm but less than 65 cm, basal length of 
skull more than 95 mm, but less than 112 mm. Color grayish- 
ocherous with gray hairs, back side of ears not black, end of tail 
not white (V.H.). 

Description 

In general construction, the corsac is a typical fox and very similar 
to the red fox, representing itself as a reduced copy. It is also 
stocky and has an elongated trunk,^ but the tail is not nearly so 
fluffy and is relatively somewhat shorter. Its length constitutes half 
the length of the trunk or slightly more; in the standing animal, it 
reaches the ground. The corsac has relatively somewhat larger 
ears; they have broad bases and pointed tips. 

Number of teats unknown. 

Winter fur dense, soft and silky. Its general color is pale-yellow 
gray with ocherous tints. Along the middle of the back, a noticeable 

Юп title page, designated 1934, on cover, 1935. Appeared in 1935 (Signed 
by press). 

''The idea that the corsac has longer limbs than the red fox (Ognev, 1931) is 
untrue. 



398 

brownish tone and well-developed "silvery", resulting from silvery- 
white ends of guard hairs. Strongly developed silvery tint sometimes 
makes fur of the animal silvery-gray on the back. Sometimes, on 
the contrary, brownish tone quite strongly developed. Shoulders 
same colored as on dorsum. Side also with same tone, but consid- 
erably lighter. Lower surface of body, including chest and inguinal 
274 region, white or with light yellowish frosting. Fore legs light-yellow 
anteriorly, laterally rusty-yellowish; posteriorly, same color but paler. 

Head grayish-ocherous or brown in color, darker on forehead. 
Around eyes color lighter, a dark field of triangular form between 
anterior corners of eyes and upper lip. Around mouth, on throat and 
along lower neck, color white with weak yellowish tint. Rear side 
of ear ocherous-gray or reddish-brown, bordered anteriorly along 
edge of ear in white, interior of ear pinna covered with yellowish- 
white hairs. 

Tail covered with dense, fluffy hairs, and has a dirty dark- 
ocherous or gray-brownish general tone. On dorsal side it has blackish 
tinge; on basal half — 6-7 cm from base in region of supracaudal 
gland, is found a dark spot. At base, brownish tone noticeable. 
Lower side pale. Tip of tail dark, usually almost black. Underfur 




Fig. 105. Corsac fox, Vulpes cor sac L. Sketch by A.N. KomaroV. 



399 






275 



Fig. 106. Skull of corsac fox, Vulpes corsac L. 



400 

ash-gray, brownish at ends or shghtly rusty. Guard hairs gray at 
base, white at tip, pale-brown in middle. Individual guard hairs have 
two dark zones with an intermediate white one. 

Summer fur differs greatly from winter — it is short, sparse and 
coarse; tail hairs are also considerably shorter. In this fur, the 
corsac fox looks wiry and long-legged, with a large head. General 
color is more even — back and sides are nearly undifferentiated, 
dirty-ocherous or dirty-sandy, dull. Gray hairs are not developed in 
summer. 

There is no sexual dimorphism in color. 

In general, the skull of the corsac fox resembles that of the red 
fox, but, irrespective of the smaller dimensions, differs from it in 
many essential characteristics. In general appearance the skull is 
much shorter and broader and not at all elongated. The general 
outlines of the zygomatic arches and rostral region are reminiscent 
of a broad wedge — zygomatic arches are very broad posteriorly, 
and the skull in the region of the infraorbital foramen is broad, and 
not as compressed as in the red fox. Therefore, the line of the 
zygomatic arches merges into the line of the rostrum more gradu- 
ally, without such a break as in the red fox. The rostral region is 
relatively short — ^the ratio between the length of the nasal bones and 
the length of the distance between their posterior end and the 
posterior-most point of the occipital region, is 1 : 1.6—1.8. 
277 The braincase protrudes behind the transverse line passing 
through the articular condyles of the lower jaw for a shorter dis- 
tance than in the red fox. The general volume of the quite convex 
and swollen braincase is relatively greater than in the red fox. It is 
comparatively short, but widened posteriorly. The line of the upper 
profile is straighter. The occipital crest is weakly developed, the 
sagittal, very poorly and 10-15 cm in length. The temporal lines 
come off of the postorbital processes almost parallel, and very 
gradually narrow posteriorly, uniting with the sagittal crest. They 
outline a wide lyriform or triangular expanse on the frontal and 
parietal bones. Orbits are relatively large — ^their greatest height is 
equal to or more than the width of the interorbital region. Paraoccipital 
processes are weaker than in the red fox — ^their ends do not reach, 
or only just reach, the level of the lower edge of the auditory 
meatus. The length of the lower jaw symphysis is 4.5—6.5 times 
shorter than its greatest length. 



402 

Canines and camassial teeth are comparatively stronger than in 
red fox. When the jaws are closed, the tip of the upper canine 
reaches the lower edge of the mandibular bone, the apex of the 
lower canine extends beyond the edge of the alveolus of the upper. 
Sometimes, as a form individual variation (age?), reduction of the 
second lower molar is observed and the dental formula becomes 

3 14 2 
I-C-P-M- = 40. 
3 14 2 

Other than size, there are no sexual differences in the skull. 

Number of chromosomes unknown. 

Only approximate body size of the corsac fox is known, since 
the data published are those of individual measurements of animals 
"in the flesh". Data on the skull dimensions are somewhat more, 
but the material is very uneven. 

Body length, 50-60 cm; tail length, 25—35 cm; height at shoulder, 
about 30 cm; length of ears, about 8 cm; length of sole of hind foot, 
about 11 cm. 

Greatest length of skull, 102.2—124.0 mm; condylobasal length, 
98— 1 18.1 mm; zygomatic width, 57.5—71.1 mm; skull height in region 
of auditory bullae, 39.0-45.7 mm. 

Sexual dimoфhism in size is almost not expressed, and appar- 
ently males average only slightly larger than females. The belief 
sometimes reported, that females are larger than males, is incorrect 
(V.H.). 

Geographic Distribution 

Flat steppes and semideserts, in part deserts of Asia and Southeast- 
ern Europe. 

Geographic Range in the Soviet Union 

This is relatively small, but constitutes a considerable part of the 
western range of the species. It occupies the steppes and deserts 
of the southeastern European part of the Soviet Union, northern 
Caucasus, Kazakhstan, Middle Asia and Trans-Baikaliya. It is di- 
vided into two parts isolated from each other, uniting outside the 
borders of the USSR. 

The main and vastest part of the range occupies Middle Asia, 
Kazakhstan and adjacent territories. In the west, the southern 



403 

borders of the distributional area of corsac fox includes the Cis- 
Caucasian steppes on the south. Beginning on the Caspian coast at 
Makhachkala, it passes along the foothills to Khasav"yurt, farther 
to Groznyi and Mozdok, and then, approximately to Stavropol' and 
from there, embraces the Egorlyk basin, and passes on to the lower 
Western Manych and to the lower Don [rivers]. 

Along the Don, which constitutes the western border, it rises 
approximately to 39° N. Lat. (somewhat north of Volgograd)^ Thence, 
278 the line of the border, following the left bank of the Volga, reaches 
northward almost to the Samarsk bend, and then ascending somewhat 
more to the north and to the east and becoming the northern border 
of the range, it encloses the extreme southern regions of the eastern 
half of the Tatar Republic (Bugul'minsk, Bavlinsk, approximately at 
54° 30' N. Lat.) and adjoining— Bashkiriya. Farther, it passes along the 
outliers of the Bashkirian steppes and along the Obshchii Syrt it turns 
shaфly to the south to the southern end of the Urals. 

Rounding the southern spurs of the Ural mountains along the 
right bank of the Ural [river], the border on the eastern slope of the 
range goes northward and passes through Verkhnaural'sk. Thence, 
going eastward through Troitsk, Shadrinsk, Tyumen', Таги, Ishim, 
Vengerovo, Barabinsk, Omsk and Lake Chany, the boundary line 
passes through the bend of the river at Kamen' on Ob'. Following 
the left bank of the [Ob'] river, it descends to Barnaul, rounds the 
Altai from the west and along the northern slope of the Zaisan 
depression, exits into China. 

In Middle Asia and Kazakhstan, the range occupies all the level 
expanses, i.e. corsac fox is absent in Tarbagatai, Dzhungarskii Alatau, 
in the entire Tien Shan (occurs in the Karatau) and Pamir- Alai and 



277 ^There is not sufficient exact information about the normal occurrence of 
corsac fox to the west of the Don. Therefore there is no basis at present to assign 
the western border of the range along the "eastern part of the Dnepr-Don 
interfluve (Pidoplichko, 1951). At the same time, it is probable that some centuries 

278 ago, when no cultivation was present in the southern European part of the Soviet 
Union and people were very scarce, the corsac fox lived somewhat farther west 
than now; probably it lived normally in the Priazov steppes. However, positive 
information on this is absent. 

It is not excluded that there were intrusions farther to the west as far as 
Zaporozhsk district and western Priazov. However, intrusions to Zaporozhsk 
district were extremely rare (Sharleman' [Charlemagne], 1937). There is no basis 
for believing that corsac fox normally liyes and is even sufficiently common (at 
Korsak-Mogila; Sharleman' [Charlemagne], 1948). 



404 

Kopet-Dag. It is found in the level parts of the right bank of the 
upper Amu-Dar'ya in Tadzhikistan, but it is, apparently, absent in 
the Fergana valley. From Turkmeniya, where the mountain range 
(Kopet-Dag) forms no obstacle, the range passes to Afghanistan 
and Iran, and from eastern Kazakhstan, through the Dzhungarsk 
Vorota [Gate] and Zaisan depression, into Dzhungariya. 

Another section inhabited by corsac fox is found in our country 
in southern Trans-Baikaliya. This region represents the northern 
outlier of the Manchurian and Mongolian part of the range. In the 
adjacent USSR, the range is separated from the already-described 
western part. The Trans-Baikal section of the range is very small 
and, in its turn, is divided into two parts. The first includes only the 
steppes along the Argun' and Onon, and the Aginsk steppe (the 
area lying to the west of the lower course of the Onon). The 
second section is still smaller and occupies the steppes along the 
Selenga and Dzhida in western Trans-Baikaliya. 

The northern border of the main range of corsac fox cannot be 
considered sufficiently defined. Beside the insufficient study of the 
corresponding territories, the reason for this is that corsac fox is 
characterized by more or less regular seasonal, and also irregular, 
migrations. Moreover, through the force of confluence of various 
ecological conditions, considerable fluctuations exist at places in its 
numbers, especially at the borders of the range. Finally, the corsac 
fox under harvest very easily yields to extermination, and slowly 
restores its numbers. Therefore, in individual years or periods it 
may, firstly, shift somewhat northward the northern border of its 
reproductive area or, secondly, there may occur locally quite long 
transgressions by stray individuals. 

At the same time, due primarily to limited information about this 
species, it is essentially impossible to firmly delimit breeding terri- 
tory, and territory of migration. The outlined northern border is, to 
a. considerable degree, determined by more or less regular, if none- 
theless rare, migrations, and the true limit of the breeding area lies, 
apparently, somewhat southward. Thus, in southern Tatar Republic, 
the corsac fox occurs only as a transient (migrating), and the actual 
[northern] limit between the Volga and Ural apparently lies at the 
279 latitude of the Samarsk bend (about 53°). The same thing applies 
to the extreme northern points where corsac fox is encountered 
between the Don and Volga. In exactly the same way, the outlined 
southern and southwestern border in the Cis-Caucasus is, to a 



405 

considerable extent determined by migrations. The area of normal 
occurrence here is restricted (or, until recently was restricted) to 
the semideserts to the north of the Terek; in the west it did not 
reach Stavropol'. 

Long migrations are known for the corsac fox from the steppes 
in the southern Ural foothills to Zilair and in Shadrinsk region (at 
56°), from the Privolga steppes westward to Hel'govk (about 50 
km to the north of Nogaisk on the shore of Berdyansk gulf), and 
to Pavlograd (east of Dnepropetrovsk). Within the range, there are 
sections where the corsac fox is absent (Ulutau, Kazakh upland) or 
they occur as transients (southern Pribalkhash) (see section on 
"Biology"). 

Geographic Range outside the Soviet Union 

This occupies the steppe part of northeastern China, in all events 
Barga (expanse between the Argun'and Great Khingans), and prob- 
ably the Manchurian plains in the Nonn' and Sungari basins; Inner 
Mongolia; the entire Mongolian Republic, except the taiga districts 
of the Kentei mountains, the wooded and mountainous districts of 
the Khangai [mountains], and the Prikosogol'sk district*; Dzhungariya 
and Kashgariya; Afghanistan (probably only northern regions); and, 
apparently, northeastern Iran. 
280 The limits of the distribution of the corsac fox in southern 
Central Asia is not known — it is possible that it extends to the 
ranges that delimit Tibetan Plateau on the north (see section on 
"Geographic Variation"). 

Therefore, as a whole, the range consists of two vast areas, 
western and eastern (central-Asiatic), with a narrow "isthmus" 
uniting them in the area of the Dzhungarsk Gate and Zaisan de- 
pression. 

In the Pleistocene, corsac fox was widely distributed in Middle 
and, apparently, Western Europe and it still lived in the Crimea in 
the Holocene^ (V.H.). 

*Area around Hobsogol Lake — Sci. Ed. 

^Geographic distribution of the corsac fox according to Dinnik, 1914; 
Ognev, 1931; Shergin and Frenkei', 1936; Sharleman', 1937, 1948; Heptner and 
Formozov, 1941; Heptner et al., 1950, V.A. Popov and Lukin, 1950; Shvarts, 
Pavlinin and Danilov, 1951; Pidoplichko, 1951; Kirikov, 1952; Yanushevich and 
Blagoveshchenskii, 1952; Afanas'ev, Bazhanov et al., 1953; Bannikov, 1954; 
Vereshchagin, 1958; Sludskii, 1959 and a series of other sources and on unpub- 
lished material of V.G. Heptner. 



406 




279 Fig. 108. Species range of corsac fox, Vulpes corsac L. V.G. Heptner. 

Geographic Variation 

The geographic variation is quite evidently manifested, although, 
apparently less than is sometimes considered (Ognev, 1935). 

Within the limits of the USSR, one may differentiate the follow- 
ing subspecies. 

1. Kazakhstan corsac fox, V. corsac Linnaeus, 1768 (syn- 
onym kalmykorum). 



407 

Relatively large form, with fluffy and dense winter fur of above 
described color, without admixture of reddish tones in winter fur. 

Greatest length of skull 103-1 1 1 mm, basal length 96-105 mm, 
zygomatic width 59-68 mm, greatest skull width 37-42 mm, skull 
height in the region of tympanic bullae 42.6-44 mm, length of upper 
tooth row 48-55 mm (less than 10 individuals; Ognev, 1931). 

Northern part of range of the species, to Cis- Altai steppes. Distri- 
butional limits in the south are not established — ^apparently, it does not 
extend farther than the latitude of the northern end of the Aral Sea. 

Not outside the USSR. 

2. Turkmenian corsac fox, V. с turcmenica Ognev, 1935. 

Considerably smaller than the preceding (smallest form of the 
species), with shorter, coarser winter fiir, of dirty-grayish color and 
reddish tones. 

There are no data on body measurements. 

Skull measurements of Turkmenian corsac fox (14 males and 
females; V.G. Heptner) are as follows (Table 30). 

Inhabits the southern part of the species range — ^plains of Middle 
Asia and Kazakhstan, probably, south of the parallel on the Aral 
Sea's northern end. The boundary with respect to the preceding 
form is not elucidated— apparently, there is a wide zone of transition 
between each other. 

Outside our borders — in northern Afghanistan and northeastern 
Iran. 
281 The corsac fox of the Volga-Ural steppes and right bank of the 
Volga apparently has some characteristics of transitional character 
between the normal form and turcmenica. They are described as 
kalmykorum, and their furs are considered as a particular Astra- 
khan type. 

Table 30. Skull measurements of Turkmenian corsac fox (in mm) 



Data 


Minimum 


Maximum 


Average 


Total length of skull 


102.2 


114.0 


105.8 


Condylobasal length 


98.4 


110.3 


103.0 


Length of nasal bone along midline 


31.7 


36.3 


33.9 


Zygomatic width 


57.5 


65.3 


60.4 


Greatest width of cranium 


40.7 


44.1 


42.5 


Height in region of tympanic bullae 


39.3 


43.2 


41.6 


Length of molar tooth row 


38.3 


42.5 


40.2 


Interorbital width 


18.4 


23.0 


20.7 



408 

3. Trans-Baikal corsac fox, V. с scorodumovi Dorogostaiskii, 
1935 (synonyn nigra). 

There are no data on body measurements. 

Skull dimensions of animals from southern Trans-Baikaliya, are 
as follows (Dorogostaiskii, 1935) (Table 31). 

Table 31. Skull measurements of Trans-Baikal corsac fox (In mm) 



Data 


Males (41 spec.) 


Females (37 spec.) 


M 


linimum 


Maximum 


Average 


Minimum 


Maximum 


Average 


Greatest length 


110.9 


124.0 


117.0 


110.2 


119.6 


114.6 


Condylobasal length 


106.0 


118.2 


113.5 


106.0 


117.0 


110.5 


Zygomatic width 


63.0 


71.1 


66.9 


63.0 


68.8 


65.2 


Width above canines 


18.4 


21.5 


19.9 


18.0 


21.5 


19.6 


Maximum width of 


42.0 


46.1 


44.2 


41.4 


47.7 


43.5 


cranium 














Length of nasal bone 


34.0 


43.0 


38.0 


35.0 


40.4 


37.0 


Height in region of 


41.2 


45.7 


43.7 


41.5 


45.0 


43.1 


tympanic bullae 















Found in Trans-Baikaliya. 

Outside the USSR, in the Mongolian People's Republic and 
northeastern China. 

To this form, by priority the name nigra Kastschenko, 1912 
should be assigned; it is based on a skin (without skull) without 
accurate locality reference, but which apparently originates from 
Trans-Baikaliya or the contiguous part of the Mongolian Republic. 
However, this name is hardly acceptable, since the author de- 
scribes an ordinary melanic, and, due to the defective skin, he is not 
even sure that it is assignable to this species. 



Separate forms of the corsac fox have not been described from 
that part of the range lying outside our territory. One may suppose 
that there, except perhaps in Kashgariya, the same subspecies live 
which are found in our country. 

In the western literature, there is a widely distributed idea (see, 
e.g., Ellerman and Morrison- Scott, 1951) that V. eckloni, described 
by Przheval'skii (1883) from northern Tibet, constitutes a subspe- 
cies of V. corsac. This idea is mistaken — V. eckloni is a synonym 
of V. ferrilata. The error is probably based on the fact that in 



409 

earlier works (in part, as indicated), Przheval'skii called this species 
the corsac fox (V.H.). 

Biology 

Population. In the steppes of the Pri-Black and Pri-Azov Seas, 
corsac fox became extinct, apparently, only in the 18th— 19th cen- 
turies. The cause for this was believed to be the ploughing of the 
virgin steppes (Pidoplichko, 1951; Topachevskii, 1956), but, appar- 
ently, its extinction took place under the influence of other 
factors — climatic deterioration (it became less continental) and 
282 others. Reduction in numbers of the corsac fox also may have been 
influenced by the extermination of the steppe marmot [Marmota 
bobac], the burrows of which it occupies particularly eagerly, and 
by massive reproduction of the red fox in cultivated fields, repre- 
senting not only its competitors but also its enemies. 

At the beginning of the current century, corsac fox was not 
rare in the Don steppes, in Ergeni [hills] and in the Astrakhan 
semideserts and south of the latitude of Volgograd (Artsibashev, 
1859; Dinnik, 1914; Kirikov, 1959). In recent years, its population 
even increased, and in Ergeni the animal is now common. 

In the Cis-Caucasus, this carnivore is common in the region of 
the Kumo-Manychsk canal and in the Kum-Terek interfluve. Dur- 
ing this century, in the latter region, its population was subjected to 
significant changes. Thus, in the beginning of this century, in the 
northern Caucasus, it was communicated (Dinnik, 1914) that: "Pre- 
viously, in all these very poorly settled places, corsac foxes lived in 
great numbers, but with settlement of the steppes and their plough- 
ing, these animals began to greatly decrease in numbers; in many 
places they even became completely extinct". In 1924—1925, corsac 
foxes again became numerous in the northeastern Cis-Caucasus. A 
single hunter caught 15—30 animals in one season (Ognev, 1931; 
Heptner and Formozov, 1941). By 1931, the population of this car- 
nivore was again strongly reduced. There were relatively many 
corsac foxes in the Cis-Caucasus in 1951 (Vereshchagin, 1959). 

East of the Volga on the Obshchii Syrt and adjacent regions 
northward to the upper Bol'shoi and Malyi Irgiz rivers, the corsac 
fox periodically appears common, since its skins are obtained by the 
hundreds (N. Severtsov, 1861; Zarudnyi, 1897; Bazhanov, 1930; 
A.A. Sludskii). It is very rarely found in the steppes of Bashkiriya, 



410 

penetrating in occasional years to the Beloretsk region and southern 
Tatariya (Popov and Lukin, 1949; Shvarts et al., 1951). At the 
southern end of the Ural mountains, the corsac fox becomes more 
common, for example, in Orenburg district, along the border with 
Kazakhstan. In the Volga-Ural interfluve, the corsac fox is again 
frequent and, for example, in the Ural'sk district in different years 
up to 5,500 individuals are caught in a season, while in Gur'evsk 
district — ^up to 1,700. To the south, on the Mangyshlak Peninsula 
and Ustyurt Plateau, it is common (up to 500 spec, caught per 
season). It is also common in Mugodzhar. 

In western Siberia, the corsac fox appears in various years in 
the northern forest-steppe zone and even on the edge of taiga 
forests, but here, its numbers are extremely low. In the southern 
half of the forest-steppe zone (Barabinsk and Kulundinsk steppes), 
the corsac fox is rare and not a permanent occupant. In the south- 
ern part of the steppe zone of western Siberia (Chelyabinsk, 
Kustanaisk, Severo-Kazakhstansk, and Pavlodarsk districts and Altai 
territory), the carnivore being described was common in the last 
two decades and in some places, it was abundant. In separate 
years, in each of these districts, from 1000 to 3000 skins were 
obtained. 

The abundance of corsac foxes in typical steppes may be judged 
by the number caught by hunters. In years with high populations of 
corsac fox, in central Kazakhstan, hunters catch 20-50 and more 
animals in a season. In 1947, in Tselinogradsk district, 37 burrows 
of corsac fox were found in an area of 30 km^ and of these 8 had 
litters, in 8 burrows there had been, apparently, litters, and 9 were 
inhabited by single animals. In 1948, in this same region, 34 burrows 
were found, of which only 4 had litters (Chirkova, 1952). It is also 
numerous in the semideserts of central Kazakhstan (Tselinogradsk 
and Karagandinsk districts), and also in the Pri-Irtysh steppe 
(Semipalatinsk district). In some years, in central Kazhakhstan, up 
to 4,500 corsac skins were obtained in a season. In the semidesert, 
the corsac fox periodically becomes very abundant. Thus, near the 
Kulanutmes river from July 29 to August 5, 1951, 27 burrows of 
this animal were found in an area of 1,000 hectares, of which 7 had 
litters in spring (A.A. Sludskii). In years of massive reproduction, 
some hunters capture up to 100-120 animals in one season. The 
283 corsac fox is abundant from time to time in the Pri-Aral'sk Karakum, 
common, especially in winter, in Betpak-Dala, occurs in the Pri- 
Chuisk Muyunkum and in northern Pri-Balkhash'. 



411 

In the sandy deserts south and east of Betpak-Dala-Muyunkum, 
Taukum and Sary-Ishikotrau the corsac fox, as a rule, does not live 
permanently, but comes from the north only in certain severe win- 
ters. This animal is extremely rare in the north and west of the 
sandy desert of Kyzylkum (Bogdanov, 1882). In the Amu-Dar'ya 
delta, it is rare (Salikhbaev, 1950). In Kara-Kalpakiya, from 1956 
to 1959, only 21 to 76 skins were prepared annually. In the Syr- 
Dar'ya valley, on the clayey plain Dar'yalyktakyr lying to its north, 
and in the sands of the Pri-Aral'sk Karakum, the corsac fox is not 
especially rare, as in the clayey sections of the Ustyurt. There also, 
on sands, for example Asmantai, they are scarce. In the Ustyurt 
and in the northern Pri-Caspian, corsac foxes in autumn and winter 
time are concentrated on the seashore and near the abandoned 
summer camps and villages, where they feed on refuse and 
rodents. 

In Turkmeniya, it is "very common" in many regions (Radde 
and Walter, 1889). Thus, it is common on the southern border of 
the Karakum and in the Badkhyz (Heptner, 1956). At the same 
time, in the Krasnovodsk region, this carnivore is scarce 
(Vinogradov, 1952). In the 1924/25 season in Turkmeniya, 11,522 
skins were tanned and in 1925/26 — 7,174 skins (Kogan, 1931). From 
1948 to 1959, the number of corsac foxes caught here ranged from 
694 (1948) to 2,298 (1956) per year. At the present time, in the 
majority of places in Turkmeniya, except remote sections of the 
sands, the corsac fox is common (Sapozhenkov, 1960; V.G. 
Heptner). In Tadzhikistan, corsac fox is rare in the flat southern 
and southwestern regions. In the period from 1951 to 1959, only 1 1 
(1952) to 63 (1959) skins were tanned per year. As mentioned 
above, corsac fox is absent in montane Tadzhikistan. 

In Uzbekistan, the corsac fox occupies in all of the plains and 
foothill desert and semidesert sections, and is common in some 
places. In the 1924/25 season, 8,039 skins were prepared in the 
republic and in 1925/26 — 5,442 (Kogan, 1931). In recent years, the 
number of animals everywhere decreased (Ishunin and Lustin, 1959), 
probably in connection with the intensive development of new lands, 
but in a series of districts, it is common at the present time. Thus, 
from 1956 to 1959, in Kashkadar'insk district, from 580 to 1,244 
corsac foxes were caught per year, in Bukhara — from 395 to 546, 
in Samarkand — from 288 to 474, and in the Tashkent — from 208 to 
336. In the Fergana valley, they are obtained singly from 1956- 



412 

1959, 1—12 skins. The corsac fox is rare in Korezmsk (16-88 skins 
per year) and Surkhandar'insk districts (68-94 skins per year). 

In Kirghiziya, in the 60' s of the previous century (N.A. 
Severtsov, 1873) the corsac fox was common not only in the Chuisk 
and Talassk valleys, but also in high mountane regions (Susamyrsk 
Plateau, Dzhumgal, the Lake Sonkul' depression, on the Aksai and 
in the lower reaches of the Chirchik, Arys, and Keles rivers). 
However this information is doubtful. In the last decade, the corsac 
fox was not found permanently in any place in Kirghiziya (Kuznetsov, 
1948). In the severe winters of 1948/1949 and 1950/1951, these 
animals appeared in the Chuisk valley and the foothills adjacent to 
it, and became fairly common there. In 1956-1959, Kirghiziya yielded 
131—137 skins of this animal. 

In southeastern Kazakhstan (southern Pri-Balkhash', Chu-Iliisk 
mountains, foothill regions of the Zailiisk and Dzhungarsk Alatau, 
Alakul'sk depression), the corsac fox was extremely rare and was 
caught singly until the 50' s of the present century. In many regions, 
it was completely unknown. Starting from 1951, populations of the 
animals here started to grow rapidly and in recent years in Alma- 
Atinsk and Dzhambulsk districts, up to 1000-1300 skins began to be 
taken annually. 

In southern Pri-Baikal the corsac is "very rare" (Selenga valley; 
Fetisov, 1949) and does not appear every year (Surmach, 1957). In 
southeastern Trans-Baikaliya in the middle of the last century, the 
corsac fox was very common in the montane steppes (Maak, 1859). 
"Along the Argun' and Onon they are a multitude" (Cherkasov, 
284 1884). Later, in connection with the ploughing of the steppes, num- 
bers of these animals strongly decreased in eastern Siberia and in 
the 20 's of the present century only 341 skins were tanned in 1925/ 
26 and in 1926/27, 333 skins. In the 30's, the number of corsac 
foxes here again grew, and up to 1 ,400 corsac began to be caught 
in a year (Podarevskii, 1936). In the last two decades, the number 
of the corsac foxes in Trans-Baikaliya has again decreased (see 
"Population"). 

Therefore, within the boundaries of USSR, the corsac fox is 
most numerous in dry steppes and semideserts (central Kazakhstan). 
In deserts of the northern and southern types (southern Kazakhstan, 
Turkmeniya, Uzbekistan) this carnivore is less numerous, but in 
places it is common and even abundant. 



413 

In the 1 8th century, in the territory of contemporary Kazakhstan, 
corsac was much more numerous than now. At that time, Kazakhs 
sent from 40 to 50 thousand skins every year to Orenburg and 
Troitsk fortresses (Rychkov, 1762). "Innumera copia" (innumer- 
able multitude) — ^thus was described the number of corsac foxes in 
Kazakhstan at the end of the 1 8th century and beginning of the 
19th by G. Fisher (1814). Later, as a result of intensive harvest and 
especially ploughing of the steppes, the populations of corsac fox 
started to decrease. They declined especially strongly after 1954 in 
connection with massive ploughing of virgin lands. 

The corsac fox is numerous in the Mongolian People's Repub- 
lic. In individual years, up to 18 (1928) and even 24 thousand skins 
(1931) were obtained there. About 90% of this quantity was ob- 
tained from the eastern plain-steppe and semidesert regions 
(Bannikov, 1954). In the province of Xinjiang (China), in Iran and 
in Afghanistan, the corsac fox is rare. In northern Afghanistan, 
however, it is quite common (V.G. Heptner). 

Habitat. In the Pri-Azov the corsac fox is restricted to the 
weakly undulating plains of the watersheds which are occupied by 
Pri-Black Sea herbaceous-fescue-feather grass and fescue-feather 
grass steppes, as in the southern chernozem. Here are innumerable 
ravines and gullies, at the heads of which the soils are salinized and 
have semidesert vegetation, covered mainly by fescue, meadow 
grass {Poa bulbosa) and desert-steppe shrubs: prostrate summer 
cypress {Kochia prostrata) and various species of sagebrush; also 
thickets of steppe shrubs — blackthorn {Prunus stepposa), spiraea, 
Russian peashrub (Caragana frutex) and others. These steppes 
were formerly inhabited by bobac marmots [M. bobac]. At the 
present time all of these steppes are almost completely destroyed, 
in connection with which the corsac fox lives there on the solon- 
chak [areas of saline soil] near the seashore (Korsak-mogila) 
[Corsac-grave], in elevated areas; and along southern slopes of 
gullies as well as in pastures. 

In the western half of the Cis-Caucasus, this carnivore lives in 
steppes very similar to those already described, while in the Pri- 
Caspian and the lower Volga, the corsac fox inhabits sagebrush-turf- 
grass steppes and semishrub sagebrush and Russian thistle desert 
(southern semidesert), and is also encountered on compacted sahds 
(Nogaisk steppe). In the Volga-Ural interfluve, the corsac fox in- 
habits the steppes, semideserts and deserts, but it is more common 



414 

in hilly semideserts with a low and scattered stands of grass; along 
the middle course of the Ural river, on the Obschii Syrt and on the 
upper Chagan river, it lives in the hilly steppes (Zarudnyi, 1894) and 
chiefly inhabits the slopes of hills scattered across the level feather 
grass-sagebrush semidesert (Kuznetsov, 1928). In the Volga-Ural 
sands, it is rarely met with and, apparently, is not permanent. 

During years of increase, it inhabits even the forest-steppe of 
western Siberia beyond the Ural mountains, with its flat relief where 
the low and narrow ridges ("crests") 2 to 4 m high, rarely 6 to 
10 m, alternate with the depressions occupied by lakes, rush-cov- 
ered floodlands, small birch forest — kolka ["pins"] — and 
solonchak. In the forest-steppe, the corsac fox mainly inhabits 
solonetz [alkaline] soils around large salt lakes with poor vegetation 
and occasionally the crests if they contain significant sections of 
285 virgin soil with low fescue steppe. It hunts mainly around water 
bodies with open shores, on solonchak, in depressions and along 
crests where little snow occurs in winter. It flees into kolka and 
rush-covered floodlands only during periods of ice crusts, where it 
searches for white hares [Lepus timidus] and willow ptarmigan 
[Lagopus lagopus] caught in snares (Sludskii, 1930). 

South of the forest-steppe, from the left bank of the Volga and 
to the Ob' [river], the corsac fox lives in herbaceous-fescue-feather 
grass steppes on ordinary and southern chernozem (Kulundinsk 
steppe), and also in turf-grass dry steppes lying more to the south. 
In the typical steppes, the corsac fox settles in the burrows of 
marmot and ground squirrels, located on the gentle slopes of 
elevated areas not far from the edges of solonchak, and in sum- 
mer it hunts mainly in places with scarce vegetation. There, where 
the steppes are ploughed, the corsac fox survives only in pastures 
and near bodies of waters; it is often obliged to live near villages. 

In winter, the corsac fox chooses places where the snow is 
blown clear by wind or is Hght, but it also appears in thickets of 
reeds near bodies of water. Thus, in the very snowy winter of 
1940/41, in Severo-Kazakhstansk district, corsac foxes were found 
in considerable numbers on the steep slopes of the great Lake 
Chagla depression, where there was little snow and it could prey 
on voles and mice. Driven by hunger, the corsac foxes were cap- 
tured in cut reeds growing around the lake, under which there was 
little snow and where they hunted the water voles [Arvicola 
terrestris] which were very numerous there. 



415 

In Kazakhstan, the corsac fox is especially numerous in desert 
steppes between the Ural and Emboi, along the Turgai and south 
of Lake Kurgal'dzhin and the Nura river. There is undulating or 
hilly relief with narrow river valleys and quite expansive depres- 
sions with fresh and salt lakes. In these regions, the corsac fox 
occupies gentle sides of valleys and slopes and is usually absent in 
melkosopochnik [region of small mounds or hills], occurring only 
around the base of the sopok [mounds]. Thus, southeast of Lake 
Kurgal'dzhin, near the Kulanutmes river, burrows were distributed 
on the upper parts of the slope — the watershed between the 
Kulanutmes and a deep narrow valley. From the river to the sum- 
mit of the slope was 2—3 km. The gentle declination of the slope 
was occupied by fescue-feather grass steppe with 30-50% of the 
soil covered with vegetation. In the upper part of the slope, old 
burrows of marmots and ground squirrels were encountered. Corsac 
foxes also used them as dens. 

The building of a burrow by corsac foxes on elevated areas (exr 
eluding melkosopochnik) is explained by the high flooding of the steppe 
rivers; the water level of which rises 10-14 m. Building their burrows 
in the higher part of knolls, the corsac fox in summer hunts along their 
slopes, and rarely appears at the bottom of ravines, or at the river or 
lake, but it usually avoids entering hilly country. In autumn and the 
beginning of winter, in the northem semidesert (Lake Kurgal'dzhin), it 
restricts itself to flat areas, overgrown with sagebrush and Russian 
thistle (B.D. Abaturov). 

After abundant snowfall, when the depth of snow cover in 
areas with tall vegetation reaches 20 to 40 cm, and it becomes 
more densely packed, while in areas with low vegetation cover (in 
sagebrush, sagebrush-fescue and sagebrush-saltbush [Atriplex сапа] 
associations), snow cover is shallow; the corsac fox prefers such 
sections with little snow or places with very packed snow. There, 
the winter burrows are also found, located along more elevated 
areas. 

Farther to the south, the corsac fox lives in true deserts of the 
northem type, inhabiting the entire Betpak-Dala. In summer, it is 
rarely encountered here. For nine years, on an itinerary extending 
for a total of ten thousand kilometers, only four burrows with litters 
were found. In Betpak-Dala, the corsac fox settles on the summits 
of knolls covered with small sagebrush and usually 200-300 m from 
a spring with water. In the last decade, corsac foxes became 



416 

common in the foothills of Zailiisk Alatau, where they inhabit hilly 
286 clayey deserts and semideserts, not long ago inhabited by yellow 
ground squirrels [Spermophilus fulvus], in the burrows of which 
the foxes construct their lairs. Here, the corsac fox finds favorable 
conditions, since food is abundant, snow in winter is not deep, and 
is often trampled down by numerous cattle herds grazing there. 

The corsac fox inhabits the ephemeral deserts of Turkmeniya, 
Tadzhikistan and Uzbekistan. Along the lower Amu-Dar'ya, it oc- 
cupies the old valleys and deltas, but everywhere it is quite rare 
(Kostin, 1956). In clayey deserts of southwestern Turkmeniya, it is 
met with everywhere, and is not rare in compact sands. In south- 
eastern Turkmeniya, the corsac fox is found in the clayey deserts 
and semideserts and along the edges of compact sands. It is com- 
mon in the southern margins of the Karakum; it often remains near 
wells. There, its burrows are found on the slopes of sandy hillocks 
inhabited by the great gerbil [Rhombomys opimus]. In Tadzhikistan 
and Uzbekistan, it inhabits desert areas on the plain and in the 












Fig. 109. Burrow of corsac fox. Betpak-Dala desert. Kazakhstan. 16 June, 
1956. Photograph by A. A. Sludskii. 



417 

foothills, old river valleys and the small sandy massifs. It is absent 
in tugai [riparian] forests (Chernyshev, 1958). 

In southern Pri-Baikal and southeastern Trans-Baikal, the corsac 
fox inhabits hilly steppes, well-supplied with mounds having flat 
summits 50-100 m high which are irregularly scattered or extending 
as low chains. The slopes of the sopok are gentle, and pass gradu- 
ally into wide depressions. In places on the hills and in the steppe, 
there are rocky exposures and outcrops. Among the rocks and near 
them, forbs and shrubs grow vigorously which are rarely encoun- 
tered in the ravines. On large areas, the grassy steppes stretch over 
sandy and sandy-gravelly soil, mainly wild calla* cattail and wild 
rye-cattail steppes. In the steppes, the grassy and sagebrush areas 
form a complicated mosaic. The region is characterized by low 
temperatures in winter (to - 50°C), shallow snow cover — on aver- 
age only 10 cm — and permafrost located at a depth of 2.0-2.6 m. 
In Trans-Baikaliya, the corsac fox inhabit the slopes of valleys 
well-supplied with tarbagan [Marmota sibirica] burrows which 
they occupy for their shelters. 
287 In the MNR [Mongolian People's Republic], the typical habitat 
is more or less level or slightly undulating dry steppes with small, 
shallow valleys with a cover of feather grass, feather grass-bistort 
or grass-Russian thistle. This carnivore is rarer in herbaceous steppes 
of the northern type, and still more rarely encountered in montane 
steppes, whence it penetrates along the river valleys. It avoids 
mountainous relief with cliffs and rock slides as well as true shrub 
deserts. 

Accordingly, throughout the entire range of the corsac fox, the 
main habitats are the semideserts and the dry steppes resembling 
them. In grassy steppes, it chooses areas with scattered, low grass 
and shrub cover. Everywhere, it avoids dense thickets and prefers 
the plains. The corsac fox is absent in areas with loose snow cover 
more than 15 cm in depth, but there, where snow is compact or 
blown away by wind, it may exist even when the average depth is 
30 to 50 cm deep. 

Food. The corsac fox is a carnivore, adapted to hunt small and 
moderate-size lagomoфhs and rodents, including young marmots 
and hares. In summer time, small birds, reptiles and insects also 
occupy a considerable place in its diet. In distinction from other 

*The Russian word "zmeevik" is used for 5 different genera, including 
Polygonum, Calla and ''tyrsd" is defined as Typha — ^Sci. Ed. 



418 

species of the family of our fauna, it rarely feeds on plant food. In 
case of insufficient prey, it eats carrion and refuse, often poorly- 
digestible. 

In connection with the wide distribution of the corsac fox, its food 
list varies, but everywhere rodents predominate. At the northern 
limits of the range, in the forest-steppe of western Siberia (Barabinsk 
steppe), this carnivore in summer preys on narrow-skulled voles 
[Micro tus gregalis] or Alticola strelzovi and sagebrush voles 
[Lagurus lagurus]; rarely does its prey extend to water voles 
[Arvicola terrestris], red-cheeked ground squirrels [Spermophilus 
erythrogenys], great jerboas [Allactaga jaculus*], common ham- 
sters [Cricetus cricetus], Dzhungarian hamsters [Cricetulus 
sungorus], larks, charadriiformes and ducks. In winter, it preys 
mainly on mouse-like rodents, but occasionally attacks white hares, 
willow ptarmigan, gray partridge [Perdix perdix], black larks 
[Melanocorypha yeltoniensis] and snow buntings [Plectrophenax 
nivalis]. Near villages, it feeds on carrion and refuse; several in- 
dividuals gathering around a large carcass (Sludskii, 1930). In the 
steppes of Kazakhstan, a basic role in the food is also played by 
mouse-like rodents (99.4%), among them the sagebrush vole (37.9%). 
Apparently, this rodent is more often eaten, since of the numerous 
remains of unidentified voles found in the feces a significant part 
288 also (48.5%) belong to it (Table 32). In this region, birds are also 
important in the food of corsac fox (35.5%), apparently wounded ones. 

In the steppes in winter, the corsac fox feeds on mouse-like 
rodents, but when they are scarce or not obtainable, it turns to other 
species of prey. Thus, in the heavy snow winter of 1940/41, in 
Severo-Kazakhstansk district, at Lake Chagla during a time of fro- 
zen snow-crust, corsac foxes hunted water voles in cut reeds, 
digging up their runways in the thawed soil, and also gray partridge 
which spent the night in shrubbery near the lake. These carnivores 
also remained near cattle burial grounds and villages, where they 
fed on carrion and refuse. 

In 890 feces of corsac fox collected in May— June 1947 and 
1948, near Tselinograd, remains of rodents were found in 95.4% of 
all samples investigated, corresponding to: birds in 17.0%, insects in 
22.1%, reptiles in 0.8%, plants remains in 2.1% and indigestible 
refuse (rags, ropes, etc.) in 0.4% (A.F. Chirkova). Among mouse- 
like rodents, the main mass consisted of sagebrush voles, while 

*Now called A. major — Sci. Ed. 



419 

287 Table 32. Results of feces analysis of corsac fox inhabiting the Kazakhstan 
steppes (Severno-Kazakhstansk district, Sovetsk region, near Lake Chagia, 
July, 1947; from burrow with litters; 169 samples; frequency of occurrence 

in %) 

Food type % 

Mammals 99.4 

Rodents 99.4 

Various voles 99.3 
Including: 

Sagebrush vole [Lagurus lagurus] 37.9 

Common vole [Microtus arvalis] 0.6 

Narrow-skulled vole [Microtus gregalis] 7.7 

Root vole [Microtus oeconomus] 4.2 

Unidentified voles 48.5 

House mouse [Mus musculus] 0.6 

Birds 35.5 

Ducks 14.8 

Insects 1. 1 

Beetles 7.1 



common, root and narrow-skulled voles, mole-vole [Ellobius] and 
wood mice [Apodemus] were rarer. The remains of white hare, 
red-cheeked ground squirrel, jerboa, shrews — ^brown-toothed [Sorex] 
and white-toothed [Crocidum] — ^were found on isolated occasions. 
In the semideserts and dry steppes of central Kazakhastan in 
the spring— summer period, corsac foxes hunted mainly intensively 
reproducing sagebrush voles (encountered in 96.1% to 98.7% of all 
samples examined). Other voles, rats, hares, steppe pika [Ochotona 
pusilla], hamsters [Cricetus] and dwarf hamsters [Cricetulus] were 
rarely met with in the feces and clearly played no notable role in 
its diet. At the same time, the role of ground squirrels and jerboas 
here was increased. Thus, e.g. at Karsakpai, the remains of yellow 
ground squirrels were found in 13.3% of the samples, while in the 
Atbasarsk region of Tselinograd district, in 7.1%, although there this 
ground squirrel lives at the northern limit of its range. In the latter 
region, the remains of intermediate ground squirrels [Spermophilus 
intermedius]* were found in 7.2% of the excrements, while those 
of jerboas — in 14.2% (KJi. Kydyrbaev). In the Basaga region, jer- 
boas were found in 10.7% of the samples (see Table 33). 

*Synonym, S. brevicauda; usually considered a subspecies of 
S. erythrogenys — Sci. Ed. 



420 



289 Table 33. Results of feces analysis of corsac fox Inhabiting the semldesert 
of central Kazakhstan (in % of number of samples) 



Region, season 
and year, number 
of samples 



Northern 
semidesert near 
Kulantumes 



Southern semidesert — 

Karsakpai, 

Kazakhstan upland, 

Basaga'' 



Food type 



Summer, Summer, Spring, Summer, Summer, 

1950 1951 1950 1953 1954 

417 1518 45 103 132 



Mammals 

Lagomorphs and rodents 
Steppe pika [Ochotona pusilla] 
Mongolian pika [O. pallasi] 
White hare [Lepus timidus] 
Yellow ground squirrel 

[Spermophilus fulvus] 
Ground squirrel [S. intermedins] 
Little ground squirrel 

[S. pygmaeus] 
Ground squirrel (unidentified) 
Steppe mouse [Sicista subtilis] 
Great jerboa [Allactaga jaculus] 
Small five-toed jerboa 

[A. e later] 
Thick-tailed three-toed jerboa 

[Stylodipus telum] 
Jerboas (unidentified) 
Common hamster 

[Cricetus cricetus] 
Eversmann's hamster 

[Cricetulus eversmanni] 
Gray hamster [C. migratorius] 
Sagebrush vole 

[Lagurus lagurus] 
Mole-vole [Ellobius] 
Water vole [Arvicola terrestris] 
Common vole 

[Microtus arvalis] 
Narrow-skulled vole 

[M. g regal is] 
House mouse [Mus miisculus] 
Striped field-mouse 

[Apodemus agrarius] 
Wood-mouse [A. sylvaticus] 
Small rodents (unidentified) 
Carnivores 



99.8 


99.4 


100,0 


100.0 


100.0 


99.8 


99.3 


100.0 


100.0 


100.0 


— 


0.1 


2.2 


4.0 


3.8 


— 


— 


— 


2.0 


1.5 



0.2 



0.5 



0.5 



1.0 



0.7 



98.7 



2.9 



0.2 



0.2 



0.4 



0.1 



96.6 

0.1 
0.1 



— 


0.4 


0.1 


0.3 


0.5 


0.5 



13.3 



— 5. 



— 2.2 — 



0.2 


— 


13.8 


0.3 


— 


— 


0.5 


— 


— 


— 


2.2 


— 


— 


— 


2.0 


4.3 





10.7 


0.1 


— 


- — 



0.1 — 



75.6 



0.7 — 



3.0 
96.1 



8.9 — 



9.7 



0.8 



97.0 



12.1 



(contd.) 



421 



(Table 33— contd.) 



Region, season 
and year, number 
of samples 



Northern 

semidesert near 

Kulantumes 

river 



Southern semidesert — 

Karsakpai, 

Kazakhstan upland, 

Basaga^ 



Food type 



Summer, Summer, Spring, Summer, Summer, 
1950 1951 1950 1953 1954 

417 1518 45 103 132 



Weasel [Mustela nivalis] 


0.5 


0.3 








Corsac fox (wool) 


— 


0.2 


— 


— 


— 


Birds 


18.7 


16.7 


— 


5.8 


6.8 


Ruddy Shelduck 


0.2 


— 


— 


— 


— 


Ducks (unidentified) 


1.2 


— 


— 


— 


— 


Large birds 


4.6 


0.7 


— 


— 


— 


White-winged lark 


2.4 


0.9 


— 


— 


— 


Small birds (unidentified) 


10.6 


16.0 


— 


— 


— 


Duck eggs 


0.2 


0.1 


— 


— 


— 


Reptiles 


1.7 


3.1 


— 


2.0 


1.5 


Sand lizard 


0.1 


1.3 


— 


— 


— 


Lizards (unidentified) 


0.5 


1.1 


— 


2.0 


1.5 


Patterned rat-snake 


— 


0.1 


— 


— 


— 


Mamushi 


— 


0.1 


— 


— 


— 


Snakes (unidentified) 


0.2 


0.5 


— 


— 


— 


Insects 


22.0 


17.4 


— 


15.8 


4.5 


Locust 


— 


1.1 


— 


— 


— 


Beetles 


22.0 


16.3 


— 


— 


— 


Eggs of praying mantis 


— 


0.1 


— 


— 


— 


Plants (grasses) 


0.2 


0.1 


— 


— 


— 


Carrion 


— 


0.1 


— 


— 


— 



''According to the data of G. Shubin. 

In places where the tarbagan marmot is still numerous, it appar- 
ently plays a considerable role in the summer diet of corsac foxes, 
since in Trans-Baikaliya, corsac foxes catch young bobaks and 
feed on the dead bodies of adult animals (see below). 

In the semidesert, the role of reptiles among the corsac fox's 
food is negligible (1.5—3.1%). The same may be said about insects 
in years of rodent abundance (see Table 33). In a time of depres- 
sion of mouse-like rodent populations (1960), in the region of Lake 
Kurgal'dzhin, the feces of corsac fox consisted, to a considerable 
degree, of remains of grasshoppers (B.A. Abaturov). Birds have a 
more basic importance in the food of the corsac fox in semideserts 
(5.8-18.7% occurrence). In summer, this carnivore almost never 
eats carrion, but of plants it consumes small quantities of grass 
leaves. 



422 

In winter, with sagebrash voles abundant, and obtainable, the 
corsac fox continues to hunt this rodent principally, and, for ex- 
ample, in the region of Lake Kurgal'dzhin in the winter of 1959/60, 
all of its feces consisted entirely of the remains of mouse-like 
rodents. Moreover, in river floodplains, they successfully attack 
white hares; hunt black larks hiding in the snow at night from bad 
weather. Holes of corsac fox were found in early spring which 
contained shells of bird eggs buried by them in the ground. In the 
winter of 1960/61 in the region of Lake Kurgal'dzhin, when sage- 
brush voles were very scarce, corsac foxes restricted themselves 
to harvested fields and roads. In fields, they hunted harvest mice 
[Micromys minutus], house mice and common voles, and also gath- 
ered the seeds of cereal crops. In winter, feces were, mainly, 
mouse-like rodents, but contained also remains of seeds of com 
[maize], oats and barley. The hungry animals dug out from under 
the snow old boots, soles, rags, ropes and gnawed them (B.A. 
290 Abaturov). There, they also picked up small fish discarded by fish- 
ermen. In Tselinograd district in the winter of 1940/41, which 
distinguished itself by its abundant snow and frozen crust, corsac 
foxes destroyed white and common hares so intensely that their 
number became considerably reduced. An analogous picture was 
also observed there in the winter of 1953/54, After a thaw and the 
formation of a strong ice crust, corsac foxes began to starve and 
appeared in great numbers in the river floodplains and clearings, 
where they intensively hunted white hares or remained near vil- 
lages where they fed on refiise. Feeding of corsac foxes in winter, 
a time of famine for them, on refuse near populated areas, and 
even the outskirts of such large cities as Pavlodar and Karaganda, 
is a common phenomenon. 

In deserts of the northern type, extending in Kazakhstan from 
the northern Pri-Caspian to the Alakul'sk depression, rodents con- 
tinue to predominate in the summer food of corsac foxes (87.3—100% 
of all occurrences) (see Tables 34 and 35), but the importance of 
different species of animals changes greatly. Mouse-like rodents, 
especially the sagebrush vole [Lagurus] stand as secondary* prey. 
Only in the foothills of the Zailiisk Alatau and in a few other similar 
places, was the social vole [Microtus socialis] encountered in 
70.8% of occurrences, and in the foothills of Bulattau (northeastern 
Betpak-Dala) — ^in 38.4% of samples studied. 

*From context, probably should read "primary" — Sci. Ed. 



423 

291 Table 34. Results of feces analysis of corsac fox inhabiting northern 
type clayey deserts in Kazakhstan (in % of sample number) 



^\^^ Region, year, num- 


Pri-Caspian 


Foothill, 


Alakul' 


^^\^ ber of samples 


lowland, 


Zailiisk 


depression. 


^\^ 


Makat 


Alatau, 
Samsy 


Uch-Aral 


^\^ 


May 


May-June 


April— June, 


Food type ^\^ 


1934 


1954-1955 


1954 


^^^ 


7988 


209^ 


13110 


Mammals 


— 


99.0 


100.0 


Lagomorphs and rodents 


— 


99.0 


100.0 


Hares [Lepus sp.]* 


LO 


— 


— 


Yellow ground squirrel [S. fulvus] 


— 


40.2 


— 


Little ground squirrel [S. pygmaeus] 


12.0 


— 


— 


Great jerboa [Allactaga jaculus] 


0.6 


0.5 


— 


Earth hare [A. Jaculus]** 


1.1 


— 


— 


Small five-toed jerboa [A. elater] 


— 


1.0 


— 


Fat-tailed jerboa [Pygerytmus platyurus] 


0.2 


— 


— 


Thick-tailed three-toed jerboa 


4.0 


— 


— ' 


[Stylodipus telum] 








Jerboas (unidentified) 


19.1 


— 


5.3 


Migratory (gray) hamster 


— 


— 


19.1 


[Cricetulus migratohus] 








Eversmann's hamster [C eversmanni] 


1.6 


— 


— 


Tamarisk gerbil [Meriones 


— 


11.5 


2.3 


tamahscinus] 








Mid-day gerbil [M. meridianus] 


0.6 


— 


— 


Gerbils (unidentified) 


3.5 


— 


— 


Mole-vole [Ellobius] 


— 


6.0 


— 


Muskrat [Ondatra zibethicus] 


— 


— 


13.0 


Social vole [Microtus socialis] 


8.2 


70.8 


5.3 


Root vole [M. oeconomus] 


— 


— 


7.3 


Voles (unidentified) 


— 


,— 


22.9 


House mouse [Mus musculus] 


— 


6.0 


1.5 


Mouse-like rodents 


22.5 


— 


29.6 


Rodents (unidentified) 


7.0 


— 


— 


Carnivores 


— 


0.5 


— 


Steppe polecat [Mustela eversmanni] 


— 


0.5 


— 


Insectivores 


1.5 


— 


— 


Long-eared hedgehog 


L5 


— 


— 


[Hemiechinus auritus] 









(contd.) 

*In the Russian original, zayats-rusak, which is the name applied to L. 
europaeus. This is evidently a lapsus, and L. tolai is meant — Sci. Ed. 

**In the Russian original, zemlyanoi zaichik, another common name for the 
great jerboa. Probably a lapsus — Sci. Ed. 



424 



(Table 34— contd.) 



^\^^ Region, year, num- 


Pri-Caspian 


Foothill, 


AlakuP 


^\^^^ ber of samples 


lowland. 


Zailiisk 


depression. 


^\^ 


Makat 


Alatau, 
Samsy 


Uch-Aral 


^^^ 


May 


May— June 


April-June, 


Food type ^\^ 


1934 


1954-1955 


1954 


^\^ 


7988 


2099 


13Г° 


Birds 


10.5 


3.0 


3.8 


Gray partridge [Perdix perdix] 


0.3 


— 


— 


Merlin [Falco cohimbarius] 


0.1 


— 


— 


Kestrel [Falco sp.] 


0.1 


— 


— 


Gray lark [Calandrella pispoletta] 


0.2 


— 


— 


Passiformes (unidentified) 


— 


3.0 


2.3 


Birds (unidentified) 


9.8 


— 


1.5 


Reptiles 


1.4 


0.5 


43.5 


Lizards 


— 


0.5 


30.5 


Snakes 


— 


— 


13.0 


Insects 


1.6 


16.0 


— 


Locust 


— 


— 


1.5 


Mole crickets 


— 


— 


38.9 


Beetles 


— 


16.0 


29.7 


Insects (unidentified) 


— 


— 


45.0 


Plants (grass leaves, seeds of 


1.5 


6.0 


6.9 


barley and wheat). 








Carrion (of sheep, camels) 


2.1 


— 


— 



«Kolosov (1935). 
^Kh. Kydyrbaev. 
^°Khusainov (1959). 



In the food of desert corsac foxes, jerboas play an important 
role. In the Pri-Caspian lowlands, they were noted in 25%, while in 
the clayey-gravelly Betpak-Dala — ^in 30.2-77.8% of examined samples. 

Rodents of medium size occupy second place. In the Pri-Caspian 
lowland, these are the little ground squirrel [S. pygmaeus] (12.0% 
occurrence), in the northwestern part of the Betpak-Dala — ^the in- 
termediate ground squirrel [S. intermedius] (43.7-61.9% occurrence) 
and the steppe pika [O. pusilld] (53.8% occurrence); lastly, in the 
foothills of the Zailiisk Alatau — the yellow ground squirrel [S.fulvus] 
(40.2% occurrence) (see Tables 34 and 35). In the southern part 
of Betpak-Dala, the great gerbil [R. opimus] was noted in 92.0% 
of feces examined, and on the edge of Taukum (southern Pri- 
Balkhash) — in 40%. In the latter region, jerboas (40%), social voles 



425 



292 Table 35. Results of feces analysis of corsac fox inhabiting desert of 
the northern type — Betpak-Dala (in % of sample number) 



^^^^^ Region, year 


Southern 


Central 


Northeastern 


^^^^^umber of samples 


(Koktal) 


(Sorbulak) 


(Bulattau) 


^'^■^--v^^ 


June 


May 


May 


May 


Food type ^^-^^^^ 


1955, 


1954, 


1956, 


1951, 


^^^^ 


112" 


103 


63 


13 


Mammals 





100 


100 


100 


Lagomorphs and rodents 


— 


96.1 


87.3 


92.3 


Steppe pika [Ochotona pusilla] 


— 


54.4 


— 


— 


Mongolian pika [O. pallasi] 


— 


— 


— 


53.8 


Tolai hare [Lepus tolai] 


4.5 


— 


— 


— 


Intermediate ground squirrel 


11.6 


43.7 


61.9 


7.7 


[S. intermedins] 










Jerboas (unidentified) 


42.0 


77.7 


30.2 


23.0 


Migratory (gray) hamster 


7.2 


1.0 


— 


— 


Great gerbil [R. opimus] 


92.0 


— 


— 


— 


Voles (unidentified) 


— 


1.0 


1.6 


38.4 


Carnivores 


— 


— 


— 


7.7 


Weasel [Mustela nivalis] 


— 


— 


— 


7.7 


Ungulates 


— 


2.9 


11.1 


— 


Saiga [Saiga tatarica] 


— 


2.9 


7.9 


— 


Goitered gazelle [Gazella 


— 


— 


3.2 


— 


subgutterosa] 










Insectivores 


1.8 


1.9 


3.2 


— 


Long-eared hedgehog [Hemiechinus 


1.8 


1.9 


3.2 


— 


auritus] 










Birds (passiformes) 


2.7 


12.6 


31.7 


7.7 


Reptiles 


7.1 


1.9 


15.9 


7.7 


Lizards 


6.2 


1.9 


7.9 


— 


Snakes 


0.9 


— 


7.9 


7.7 


Insects 


34.0 


19.4 


36.5 


15.4 


Beetles 


34.0 


19.4 


36.5 


15.4 



(20%) and insects (beetles) (40%) of feces studied are of greatest 
importance in food of the corsac fox. 

Near a burrow with a Htter (14 May 1952, at the edge of 
Taukum desert), remains of black-bellied sandgrouse and steppe 
larks were found, and at a family burrow in the Hi' delta (May 
1961), remains of house mice, narrow-skulled voles [M gregalis], 
muskrats and Semirechensk pheasants were found. The presence 
292 of remains of saiga and goitered gazelle in corsac fox feces in 



"M.l. Ismagilov. 



426 

spring must be attributed to carrion feeding. In some deserts birds, 
mainly passeriformes, sometimes occupy an essential place for the 
described predator (up to 3 1 .7% occurrence) (see Table 35). Corsac 
fox attacks even such large birds as graylag goose [Anser anser]. 
At the end of July 1936, on Lake Telikul', was observed how it 
concealed itself at midday from adult birds sitting on the shore. It 
successfully attacked young wild geese and ducks on Barsa-Kel'mes 
Island (Aral Sea; Sludskii, 1939). 

The corsac fox readily feeds on lizards, snakes (eastern snake, 
chicken and rat snakes) and young steppe [Horsfield's] terrapins 
especially in years with low numbers of rodents. In the Alakul' 
depression, in spring and summer 1954, when the number of ro- 
dents was low, 43.5% of corsac feces near burrows with litters 
contained remains of reptiles, mainly lizards (30.5%). Insects are 
regularly eaten by corsac foxes in the desert and in considerable 
quantities, but usually do not play an important role in the diet. 
During insufficiency of rodents, they occupied a prominent place in 
the nutrition of a corsac fox family in Alakul' depression in 1954 
(see Table 34). Mole crickets (38.9% occurrence) and beetles are 
particularly often eaten. Plant remains are rarely found. Usually, 
these are grass leaves, and in the foothills of the Zailiisk Alatau, the 
animals in spring gleaned seeds of wheat and barley remaining on 
the soil surface after sowing. 

When it turns cold and the snow falls in the desert and semi- 
desert zone, the corsac fox is deprived of many foods: rodents 
293 entering into winter tOфer, migratory birds, reptiles and insects. The 
remaining voles, pikas and gerbils which retain an active mode of 
life exhibit reduced surface activity or shift to a subnival mode of 
life. Therefore, the majority of corsac foxes leave summer habitats 
and migrate in search of food to the river valleys (Ural, Emba, Syr- 
Dar'ya, Chu, Hi), and to sand [deserts] (Pri-Aral Karakam, Pri-Chu 
Muyunkum, Sary-Ishikotrau), and sometimes northward into the dry 
steppes. In winter both migratory and sedentary animals feed mainly 
on the remains of saigas and goitered gazelles preyed on by wolves, 
and on carrion. In searching for carrion and refuse, corsac foxes 
move at that time nearer to populated places and roadbeds of 
railway lines. Feeding on the remains of the wolfs prey, corsac 
foxes become their true commensals. For instance, in the Ural- 
Kushum interfluve, in the winter of 1953/54, in December— January, 
corsac foxes fed mainly on the carcasses of male saigas killed by 



427 

wolves. In 10 animals, caught in the Betpak-Dala desert from 
November 27 to December 21, the remains of saigas were found 
in 70% of the stomachs. In the middle of December, 1955, in the 
central part of Betpak-Dala, several corsac foxes were found at 
each saiga carcass. 

Food of the corsac fox in southern (ephemeral) deserts has not 
been studied. Judging by the faunal composition of these deserts, 
one may suppose that in summer it feeds there, apparently, on great 
and red-tailed [Libyan] gerbils, yellow ground squirrel, various 
species of jerboas and birds, reptiles and insects, and in winter, on 
gerbils and rarely on jerboas and birds. 

In the steppes of southeastern Trans-Baikaliya summer food of 
the corsac fox consists mainly of rodents (99.2—100% occurrence) 
(see Table 36). It quite often feeds on tarbagan marmots (18.8— 
71.0% occurrence) and Daurian pika (29.2 to 62.8% occurrence). 
Voles and hamsters stand in second place in its food, then birds, 
and lastly — ^tolai hare, Daurian ground squirrel, jerboa [Allactaga 
sibirica], insectivorous and carnivorous mammals, and insects. Char- 
acteristically, in Trans-Baikaliya, reptiles, which are rare there, are 
lacking in the food of corsac fox. It more often feeds on plant than, 
for example, in the steppes of Kazakhstan. 

The corsac fox, attacking a tarbagan marmot, causes relatively 
little harm, except to the young. These rodents do not especially 
fear corsac foxes and rarely conceal themselves in burrows on 
their approach. They only interrupt feeding, sit "like a stump", and 
call. Not rarely one may observe how an adult tarbagan marmot 
attacks a corsac fox, jumping out from its burrow or drawing nearer 
and nearer, and drives it away from the mound. Tarbagan marmots 
a year old or more, on being kept in a cage with an adult corsac 
fox, remained alive during the course of several days, notwithstand- 
ing that the carnivores were not fed during this time. Under similar 
conditions, young tarbagan marmots (several months in age) quickly 
became victims of the corsac foxes (Brom, 1952). Remains of 23 
tarbagan marmots, found near and in the burrow of a corsac fox 
consisted of: juveniles (mendef) — 4, animals in their second year 
(koteP) — 16; and adults — 3 (Nasimovich, 1951). It is believed that the 
corsac fox can kill only young tarbagan marmots, but among the 
adults, it takes solely dead ones (Nasimovich, 1951; Brom, 1952). 
Sometimes marmot families with young live next to a corsac fox 
litter and the carnivores do not touch them. 



428 



294 Table 36. Food of corsac foxes in the steppes of southeastern Trans- 
Baikal based on analysis of samples from contents of their stomachs 
and feces (in % of sample numbers) 



^"\_^ Season of 




Summer 




Winter 


^\..^^ year, material 


52 


feces 


38 


feces 


^Ss,,^^^^ 








^\^^^ stomachs^^ 


56 


500 


sto- 


323 


Food type ^\,^^^ 




samples^^ 


sam- 
ples^3 


machs^'' 


samples 


Mammals 


100.00 


100.00 


99.4 


52.6 





Ьа§отофЬ8 and rodents 


100.00 


100.00 


99.2 


39.4 


94.2 


Tolai hare [Lepus tolai] 


— 


1.5 


— 


7.9 




Daurian pika [Ochotona 


32.6 


29.2 


62.8 


— 


51.1 


daurica] 












Daurian ground squirrel 


3.8 


7.7 


1.0 


— 


— 


[Spermophilus dauricus] 












Tarbagan marmot [Marmota 


71.0 


60.0 


18.8 


5.2 


50.8 


sibirica] 












Jumping jerboa 


1.9 


6.1 


2.0 


2.6 


0.3 


[Allactaga sibirica saltator] 












Daurian [striped] hamster 


— 


— 


■ — 


10.5 


2.5 


[Cricetulus barabensis] 












Dzhungarian hamster 


7.7 


1.5 


■ — 


2.6 


2.8 


[Phodopus sungorus] 












Hamsters (unidentified) 


7.7 


12.3 


0.6 


— 


6.8 


Trans-Baikal tsokor [Л/уо5/?а/ш: 


] - 


— 


— 


2.6 


— 


Michnoi vole [Microtus michnoi] — 


— 


— 


5.2 


— 


Narrow-skulled vole 


7.7 


9.2 


— 


2.6 


3.7 


[M. gregalis] 












Brandt's vole [M. brandti] 


9.6 


6.1 


— 


7.9 


5.6 


Voles (unidentified) 


1.9 


12.3 


13.0 


21.0 


12.7 


House mouse [Mus musculus] 


— 


1.5 


— 


— 


0.3 


Murids (unidentified) 


5.8 


3.0 


16.8 


— 


— 


Chiroptera 


— 


1.5 


— 


— 


— 


Insectivores 


1.9 


3.0 


— 


7.9 


1.2 


Daurian hedgehog [Mesechinus 


— 


3.0 


— 


2.6 


1.2 


dauuricus] 












Red-toothed shrews [Sorex sp 


.] 1.9 


— 


— 


5.2 


— 


Carnivores 


7.7 


1.5 


0.2 


5.2 


1.2 


Steppe polecat [Mustela 


— 


— 


— 


5.2 


— 


eversmanni] 












Solongoi [M. altaica] 


— 


— 


0.2 


— 


— 


Corsac fox 


7.7 


1.5 


— 


— 


1.2 


Ungulates 


3.8 


15.6 


0.2 


2.6 


3.4 


Mongolian gazelle [Procapra 


— 


15.6 


— 


— 


3.4 


gutterosa] 













{contd.) 



\ 



(Table 36— contd.) 



429 



^4. Season of 




Summer 




Winter 


^4. year, material 


52 


feces 


38 


feces 


^s^ 








^"^ stomachs'' 


- 56 


500 


sto- 


323 


Food type ^\^ 




samples'^ 


sam- 
ples'3 


machs'" 


samples 


Roe deer [Capreolus capreolus] 


— 


— 


— 


2.6 


— 


Carrion (domestic animals) 


3.8 


27.6 


0.2 


23.7 


32.7 


Birds 


7.7 


15.3 


33.4 


26.3 


13.0 


Daurian partridge [Perdix 


— 


— 


— 


7.9 


— 


dauurica] 












Mongolian lark 


— 


— 


— 


2.6 


— 


[Melanocorypha mongolica] 












Small birds 


— 


— 


— 


10.5 


— 


Insects 


55.7 


27.6 


9.8 


2.6 


3.4 


Plants 


32.7 


37.0 


1.8 


10.5 


30.6 


Refuse (pieces of twine, rags. 


— 


3.0 


— 


59.9 


9.3 


dressed skin, etc.) 












Soil, pebbles 


63.4 


63.0 


— 


— 


13.3 



'2Brom(1952). 
'^Nasimovich (1951). 
'"Skalon(1936). 

The negligible percent occurrence of mouse-like rodents, espe- 
cially Brandt's vole, in stomachs and feces, is probably explained 
by the small numbers of voles present in the years when the ma- 
terials were collected. Occurrences in summer feces of Mongolian 
gazelle and domestic animals may be inteфreted as the remains of 
wolf meals and carrion. Observations exist of corsac foxes catch- 
ing fish in the summer of 1 944, in the dried-up lakes on the floodplain 
of the Argun' river (Brom, 1952). In half of all the examined 
stomachs of corsac foxes and in 25% of their feces are found 
insects, though as separate individuals. Of plant food, fruits of 
sweetbrier were found. 

In Trans-Baikaliya, the winter ration of the corsac fox hardly 
differs from summer (see Table 36). Only in individual years, dur- 
ing low rodent populations, feeding on carrion and various refuse 
grows. In winter, a great quantity of tarbagan marmots are found 
in stomachs and feces of corsac foxes (up to 51.1% occurrence). 
Apparently, at this time, corsac foxes feed on carcasses of tarbagan 
marmots, which die in summer burrows and were preserved there 
by the cold (Cherkasov, 1884; Brom, 1952). Through tracks, it was 
observed how the corsac foxes dig out from the snow the 



430 

entrances to abandoned marmot burrows and enter them. Frequent 
295 feeding by corsac foxes on carcasses of marmots indicates that in 
these regions, intensive epizootics occur among rodents. Occur- 
rences of remains of Mongolian gazelle and roe deer are explained 
in that the corsac fox in this region, as in Kazakhstan, feeds on the 
remains of wolf prey and on carrion. 

In the Mongolian People's Republic, Brandt's vole constitutes 
the food base of the corsac fox in summer and winter, while other 
rodents (gerbillines and hamsters) are of secondary importance. 
Thus, on May 24, in the Choibalsan region in a burrow with a litter, 
91 carcasses of mammals were found, of which 80 were Brandt's 
voles. In a burrow found on July 9 near Ulan-Tsirik, among 46 
animal carcasses, 45 belonged to this same vole (Kucheruk and 
Dunaeva, 1948). In Kentei aimak*, in two corsac fox burrows 
excavated in May and June, remains of Mongolian gazelles, Cen- 
tral-Asiatic [upland] buzzard [Buteo hemilasius] and horned lark 
[Eremophila alpestris] were found, and in feces, the remains of 
Brandt's vole and birds. Brandt's vole, within the limits of its range, 
constitutes the main food of the corsac fox (Skalon, 1949). In the 
stomachs of corsac foxes caught in spring and the beginning of 
summer, Brandt's vole, Daurian pika, quite a few orthopterous 
insects and bones of small fish were present. The corsac fox eats 
selitryanka fruits {Nitraria schoberi). More or fewer numbers of 
remains of orthopterans may also be found in winter feces of the 
corsac fox. Hunters believe that grasshoppers are a winter tidbit 
for this carnivore; therefore, they already prepare them early in 
autumn and use them in quantity as bait in traps. In southwestern 
Khangai, in winter, corsac foxes also feed mainly on Brandt's 
voles, but also eat insects. Corsac foxes, especially young, feed on 
orthopterans in other seasons also; with an abundance of locust, the 
stomachs of the carnivores are literally stuffed with these insects 
even during times of mass reproduction in Brandt's voles (P.P. 
Tarasov). 

In the Mongolian Republic in winter, corsac foxes often feed on 
carrion and refuse and closely approach populated places, appear- 
ing even at the outskirts of Ulan-Bator [Ulaanbaator]. Sometimes, 
corsac foxes become commensals with respect to diurnal raptorial 
birds — ^upland buzzard {Buteo hemilasius) and saker falcon (Falco 
cherrug milvipes), many of which winter in the northern part of 

*Province — Sci. Ed. 



431 

the Mongolian Republic. Corsac foxes gather remains of their prey, 
and at this time the stomachs of the animals become filled with hind 
limbs, intestines and heads of voles; i.e., all the parts which 
buzzards and falcons leave rather than eat (P.P. Tarasov). Feeding 
mainly on rodents, corsac foxes in the steppes and semideserts are 
concentrated in places with an abundance of sagebmsh voles or 
Brandt's voles, migrating there from the neighboring regions 
(Nekipelov, 1935; Brom, 1952; Peshkov, 1954). They also gather in 
regions where epizootics are spreading among rodents, or where 
mass mortality has occurred among wild ungulates and cattle during 
a prolonged period of continuous snow cover [dzhui]. 

With insufficiency of mouse-like and other rodents, or when 
they are unavailable, corsac foxes suffer from hunger, markedly 
lose weight and even die. In search of prey, they undertake migra- 
tions of considerable distances (see later). Gorsac foxes may remain 
without food for 7-15 days and maintain their activity. During the 
course of such a period, they sometimes do not leave the burrow, 
particularly if a trap is set at its entrance. 

Corsac foxes held in captivity in autumn consumed the follow- 
ing amounts of food: males — from 120 to 480 g, averaging 218—347 
g (various individuals); females — ^from 100 to 395 g, averaging 245— 
259 g (two individuals; Brom, 1952). In Betpak-Dala, in the animals 
caught in December in the early morning hours or at night, the 
weight of the contents of a fully filled stomach was equal to 100- 
130 g. Taking into consideration that the corsac fox hunts twice 
daily — at dawn and at night — its natural daily ration in the wild is 
approximately equal to 200-260 g, comparable to the contents in 
captivity. In ofder to be sated, it is sufficient to it to eat one pika 
(200 g) or about five voles. 
296 In Betpak-Dala, in individual stomachs, the following foods were 
found: female, on May 17 — intermediate ground squirrel (adult), 
darkling beetles; subadult male, on October 8— ^ay lark [Calandrella 
pispoletta], mole crickets (7); female, on October 18 — ^red-tailed 
gerbil [Meriones erythrourus], eastern sand snake; female, on 
November 27 — saiga meat and fat, social vole [Microtus socialis] 
(2); female on December 21 — saiga meat, red-tailed gerbil, lizard 
[Phrynocephalus sp.]. In Trans-Baikaliya, in individual stomachs 
of corsac foxes caught in January were found: 1) Michnoi vole 
[Microtus michnoi], Daurian hamster [Cricetulus barabensis], 
Daurian partridge, 10 pieces of linen rags, 2) Daurian hedgehog 



432 

[Mesechinus dauuricus], Daurian partridge, a piece of shoe, 
3) gray [brown] rat [Rattus noi'vegicus], a vole, a hamster, a 
shrew, and grass leaves (Skalon, 1936). 

With abundant prey, many carcasses may be brought into or 
near the burrow by corsac foxes (see above). In the foothills of the 
Zailiisk Alatau, carcasses of yellow ground squirrels {S. fulvus) 
were found at the burrows, but rarely other food. On 29 May 1955, 
at one burrow, 8 carcasses of yellow ground squirrels plus 8 heads 
and 4 tails, a tail of a small five-toed jerboa [A. elater] and the 
body of a lark, were found; at a second burrow, on 9 July 1955, 15 
carcasses and 8 heads of ground squirrels were found (Kh. 
Kydyrbaev). Such accumulations of carcasses are usually known 
as "reserve", but actually are the remains of collected food. Some- 
times, in periods of famine, corsac foxes eat the carcasses of their 
fellows (Brom, 1952). 

In the desert, the corsac fox is more often encountered near 
wells and springs with water, but they are not attracted to them by 
the water itself, but by the abundance of food. Even P.S. Pallas 
(1811) noticed that the corsac fox rarely drinks and can remain for 
a long time without it. "In captivity, and when fed meat, corsac 
foxes can do well without water, and if it is offered to them in the 
cage, they are completely unattracted to it" (Nasimovich, 1951). 

Home range. The dimensions of denning and hunting areas of 
the corsac fox are little known. In the Pri-Caspian lowland, when 
corsac fox was common, one litter occupied an area of 30-40 km^ 
Burrows were unevenly distributed around the territory, occupying 
the more elevated parts (Kolosov, 1935). In the steppes of Tselinograd 
district, one burrow occupied an area of 0.8 km\ and 1.9-3.7 km^ 
for one litter (Chirkova, 1952). In the same district, near the 
Kulanutmes river (semidesert), in 1951, when corsac foxes were 
common, there were 27 burrows in an area of 10 km-, 7 of which 
had litters. In the following year, after a sharp decrease in the 
corsac fox population, in the same place in an area of 2.7 km^ only 
one of 13 burrows found was inhabited; one corsac fox lived in it. 
In the Kurgal'dzhin Lake region, in the winter of 1959/60, at the 
end of February in an area of 1 km^ 5 inhabited burrows of corsac 
foxes were found, and 6 of these animals also were observed there 
(B.D. Abaturov). 

In southeastern Trans-Baikaliya, in the first half of August, the 
young of one litter lived within a radius of 300 m of the den burrow 



433 

and spent the daytime in one or another tarbagan marmot burrows. 
In this area, the corsac pups played, ran from one marmot mound 
to another and entered the occupied and unoccupied tarbagan bur- 
rows. In these same steppes, corsac foxes bringing prey to the 
pups were observed at a distance of 6 km from their den burrow 
(Brom et al., 1948). In the latter case, the hunting region of the 
corsac fox family must have occupied about 113 km^ In winter, in 
1948, in Trans-Baikaliya one corsac fox occupied 3 km- (Brom, 
1952). 

Burrows and shelters. The corsac fox is a relatively weak 
animal, has many enemies, and therefore, in open landscapes, it 
must hide in burrows and other shelters. The corsac fox uses bur- 
rows also as protection from low or very high temperatures. The 
corsac fox is a poor burrower, and cannot dig in hard ground. 
Therefore, it adapted to living in foreign burrows: of marmots — 
bobak [M. bobak] and tarbagan [M sibirica] and large ground 
squirrels — ^yellow [S. fulvus] — ^reddish [S. major] and red-cheeked 
[S. erythrogenys]. It easily enters the openings into marmot bur- 
rows, but has to widen those of ground squirrels in order to dwell 
there. Rarely, it occupies abandoned burrows of red foxes and 
badgers; only in soft sand or clay-alkaline ground does it dig itself 
297 simple and usually shallow burrows. Thus, in the Barabinsk forest- 
steppe, living places of this animal are simple in construction. They 
have 1 to 4 entrances not more than 1 meter deep. In the center 
of passages or at their end, the den chamber is built, which has no 
bedding throughout the whole year. In the forest-steppes, the corsac 
fox adopts for its own shelter the burrows of red-cheeked ground 
squirrels, widening them, or inhabits old burrows of red foxes and 
badgers. There, it digs itself only a shallow shelter with a short 
passage, known by hunters as ''dnevkf [day rest]. They use these 
only in summer for resting and for hiding in case of danger. In warm 
times of year, it usually rests by lying under a shrub, or in grass. In 
winter a corsac fox, caught by a snowstorm in the steppe, far from 
burrows, lies in a hollow dug into a snowdrift and allows itself to be 
drifted over. It rarely hides in straw- or haystacks. During snow- 
falls, it does not leave the burrow for 2-3 days (Sludskii, 1930). 

In the steppes of Severo-Kazakhstansk district, the corsac fox 
is adapted to the burrows of ground squirrels or marmots. The 
burrow depth here is 40-150 cm, being explained by the ground 
water level and deep winter snow cover. In the Sovetsk region, 1.5 



434 

km from Lake Chagla, in sagebrush-grass steppe, the den burrow 
of the corsac fox has 2 exits, the general length of the passage is 
4 m with side branches of 70 and 200 cm. Maximum depth of 
hiding passages is 40 cm. Dimensions of den chamber are 30 x 40 
cm; in it bedding is absent. On the surface between entrances, the 
animals clear a small area, known by hunters as a ''toehold'. 

In the semidesert of central Kazakhstan near the Kulanutmes 
river, the majority of 38 burrows found were situated on the sum- 
mits of ridges, part on gentle slopes and a small number in the 
bottoms of valleys. Entrances to burrows are often oriented toward 
the north-northeast or northwest, and only in individual cases to- 
ward the southeast and other directions. Entrance diameter varies 
from 20 to 30 cm, with the average equal to 22 cm. Thirty burrows 
(79%) had only one entrance, 2 had two entrances, 4 had three and 
only 2 had 4. The majority of burrows are simple, and have only 
one passage ending in a chamber. Complex burrows, with several 
entrances and side-burrows, were rarely met with. Passage length 
ranged from 140 to 450 cm, on average equaling 240 cm (of 12 
298 burrows). Greatest depth of litter in passages varied from 55 to 115 




297 Fig. 110. Corsac fox burrow under rock outcrop in semidesert at Kulanutme 
river, central Kazakhstan. 27 July 1951. Photograph by E.I. Strautman. 



435 

cm, the average equaling 79 cm (of 1 1 burrows). The upper portion 
of the burrow passages went through soft loamy soil, the chamber 
was always located in very hard gypsum of a clay layer, which 
could be cut only by an axe. All burrows were constructed in old 
ground squirrel shelters. Chambers had dimensions of 40 x 30 cm. 
In none of these was bedding found at the beginning of August. 

Soil thrown out by the animals fi^om the burrows does not form 
a high mound of elongated form in front of it as is observed at red 
fox burrows, but is scattered evenly around the entrance. Thanks 
to this characteristic, to detect a corsac burrow it is more difficult 
than that of a red fox. Around the den burrow occur many feces 
of both adults, and especially, young. Moreover, some meters from 
the entrance, in specially made pits are 2-4 "latrines." Near and 
to the side of many burrows there are 1^ ''tocholC\ on which the 
young pups played. In three cases, the living places were distrib- 
uted in groups of 3 to 4 burrows each, and each one of the shelters 
was a distance of 40 to 500 m from the other; they were connected 
to each other by trails. Such groups of burrows belong to one 
family. In the Kurgal'dzhin Lake region, in the river valley, all 
corsac fox burrows also were located in groups on elevations which 
are not submerged by snow melt-water. In other sections there, 
although single burrows were encountered, most areas were unin- 
habited (B.D. Abaturov). 

At 100 to 300 m from the den burrow, temporary burrows 
("dnevki") were sometimes met, with a passage length from 50 to 
120 cm containing no chambers. At the beginning of August, in 12 
burrows and 4 ''dnevkC\ 7 young animals were found; in one 
burrow — 2, while in the remaining — singles. Usually, 70 to 80% of 
the burrows are uninhabited. 

In semidesert, the corsac fox utilizes burrows throughout the 
whole winter. The entrance into the burrow is always cleared of 
snow, which forms a small hillock. At the beginning of winter, often 
299 there are several entrances cleared of snow, but by midwinter the 
corsac fox cleans only one entrance of snow. A burrow dug in the 
Kurgal'dzhin Lake region had, in the middle of winter, a length of 
2.5 m and depth of 1.5 m. To a distance of 40 cm, the passage was 
covered by hoar-frost, but beyond that thawed soil was found (B.D. 
Abaturov). 

In Betpak-Dala, corsac fox builds more complicated burrows 
than in the steppe and semidesert, in relation to shallow snow cover 



436 



■ S^ 



■■i'-Jiv.:''" % 

^^*-"*•■:■(■-' 



298 Fig. 111. Old den burrow of corsac fox with 23 exits. Edge of Karakum 
desert near Tashkepri on the Murgab, Turkmeniya, 23 April 1962. 
Photograph by A. A. Sludskii. 

in winter, high air temperature in summer, and loose ground which 
is easily excavated. Here, burrows are located on gentle slopes of 
ridges or on level plateaus. Of 6 burrows, one had 3 entrances, 
two — 6, the remainder — 8, 10 and 14 entrances. A family with 5 
young lived in a burrow with three entrances on level ground among 
peanut bushes. A second burrow was dug on a gentle south slope 
covered by sagebrush and peanuts near a small solonchak 300 m 
from a spring with water. It possessed 14 entrances, distributed 
over an area 7 m in diameter. The passages led to a den chamber 
located at a depth of 120 cm. At a distance of 70 m was located 
another burrow with 6 entrances and an area strongly beaten down 
by the playing of the young pups. The family of these animals used 
two burrows simultaneously, in that in the burrow with 14 entrances 
the parental pair lived, and in the burrow with six entrances — 8 
young. 

A group of three burrows was located on a slightly undulating 
clayey plain covered with sagebrush. The first possessed 18 



437 

entrances and was uninhabited. Forty meter from it was located a 
second burrow with 6 entrances used only periodically. At 20 m 
from the latter, a third, with 10 entrances, was inhabited. In it at a 
depth of 125 cm was located a den chamber, in which was caught 
an adult female and 3 young animals. Near the burrow there were 
two cleared ''toehold'. In the passages of the burrow were found 
carcasses of five intermediate ground squirrels, badly decomposed 
and with masses of fly maggots, remains of a small jerboa [A. 
elater], larks and desert wheatears [Oenanthe deserti]. At the 
entrance to the burrow, little excrement was found, but the remains 
of three intermediate ground squirrels, two eastern sand snakes and 
some larks and desert wheatears were found. In a small side- 
burrow, 12 scoфions were found. A complicated burrow with 6 
entrances was found in a solonchak on the edge of the Taukum 
sands. 

Burrows of corsac fox in the northern Caucasus are also situ- 
ated in groups (Dinnik, 1914). In this region, with sandy subsoil, it 
is easy for the animals to dig burrows, and they are built very 
widely. Each complex burrow with many deep-lying passages oc- 
cupied an area having a diameter of perhaps about 40 to 60 m in 
diameter. Sometimes, such burrows were located near each other. 
These complex burrows received the name "corsac city" from 
hunters. They believed that they were inhabited by many animals 
that lived in a sort of colony. If such vast burrows actually existed, 
they were inhabited, as a rule, by one family consisting of a pair of 
adult animals and their litter. 

On Barsa-Kel'mes Island in the Aral Sea, corsac foxes lived 
in the burrows of yellow ground squirrels, suitably widened by 
them. The length of their passages reached 4 m and were situated 
at a depth of 1-1.5 m (Sludskii, 1939). 

In the steppes of southeastern Trans-Baikaliya and the northern 
part of the Mongolian Republic, the corsac fox lives almost exclu- 
sively in the abandoned burrows of tarbagan marmots (Cherkasov, 
1884; Nekipelov, 1935, 1952; Brom et al., 1948; Nasimovich, 1951; 
P.P. Tarasov). The corsac fox does not widen the tarbagan 's bur- 
rows. Corsac foxes often visit occupied burrows of tarbagans (Brom 
et al., 1948). 

In southeastern Trans-Baikaliya, in the course of six warm 
months, the average caught out of each uninhabited [tarbagan] 
burrow was: 0.6 corsac fox in 1947, 0.2 in 1948, 0.1 in 1949 



438 

(Nekipelov, 1952). In the Mongolian Republic, the corsac fox some- 
times lives in burrows of the Daurian ground squirrel [Spermophilus 
dauricus] after widening them. Five burrows found here had from 
1 to 5 entrances. A denning burrow had one entrance and a den 
chamber of 40 x 30 cm, without bedding (Skalon, 1949). 
300 Therefore, in clayey steppes and semideserts, burrows of the 
corsac fox are relatively simple and usually not deep. In snowless 
deserts, especially in areas with soft ground, burrows occur that are 
especially complex, and den chambers are situated deep. Den cham- 
bers are also located deep in southeastern Trans-Baikaliya and the 
Mongolian Republic, where winters are very severe and snow is 
scant. The corsac fox does not strew grass in the den chamber. 
Everywhere, corsac foxes build their burrows on plains and on 
gentle slopes of ridges and hills. Burrows do not occur in ravines, 
on steep slopes or among thickets. 

Many symbionts use burrows of the corsac fox. In inhabited 
burrows, parasites (fleas) and nidicoles (flies, beetles and others)* 
are numerous. Scoфions were also found. The shelduck (Tadorna 
tadorna) and ruddy shelduck {Tadorna ferruginea) nest in aban- 
don burrows. Renard's viper (Vipera ursini) and other reptiles 
were also encountered there. 

Daily activity and behavior. The corsac fox was previously 
considered an exclusively nocturnal animal. However, recent ob- 
servations have shown that it mainly leads a crepuscular mode of 
life, and diurnal activity occurs, especially during the period the 
young are being fed, when food is insufficient, and in times of 
winter frosts. 

Due to its relatively imperfect thermoregulation, the corsac fox 
is obliged to use burrows more often than other foxes to reduce its 
activity. At times of strong freezes and wind, it remains in the 
burrow for longer than does the red fox. In the Barabinsk steppe, 
during strong snowstorms, corsac foxes did not leave the burrows 
for 2—3 days. Adapting to cold, the whole litter often lives in the 
burrow in winter. Because of this, 2 to 5 animals are successftilly 
dug out simultaneously from one burrow (Sludskii, 1930). Even in 
severe freezes, the corsac foxes, being hungry must remain long 
outside the burrow searching for prey. On calm fi-osty days, the 
animal may be observed lying on the snow near the entrance to a 
burrow, while the snow beneath it does not thaw. 

*Nest inhabitants — Sci. Ed. 




439 



^^: ■ ■■'■♦•■ 



~"ЗК 






Ifcsf*; 



300- Fig. 1 12. Corsac fox burrow near Imam-Baba. Edge of Karakum, Murgab. 
301 22 April 1962. Photograph by A. A. Sludskii. 



440 

301 The corsac fox inhabits semideserts and deserts, where air 
temperature in summer rises to 40°C and even to 50°C, and the soil 
surface becomes heated to 75°C and more. The corsac fox es- 
capes from such high temperatures in burrows and in the warm 
season, it chiefly leads a crepuscular mode of life. 

The corsac fox is a carnivore, capable of capturing food only 
in those places where there are no dense thickets (in summer) or 
deep snow cover (in winter). The corsac fox is not adapted to 
movement on snow. In the animals from Betpak-Dala, the length of 
the forelimb from the chest to the ground ranges, in males, from 17 
to 23 cm (average — 19.0 cm; 6 specimens) and, in females, from 16 
to 19 cm (average — 17.3 cm; 9 specimens), while in the Karagan 
red foxes obtained from the same region, corresponding values are 
26.3 and 24.0 cm, respectively. In spite of low weight (2.0-2.5 kg), 
the corsac fox has a great weight loading on the supporting surface 
of the foot — 61 g/cm^ for male animals from Trans-Baikaliya 
(Nasimovich, 1951). In animals from Betpak-Dala (males), it is 
from 68.3 to 80.0, or an average of 75.2 g/cm^ (4 specimens), and 
in females it is from 43.6 to 74.0 — ^an average of 54.4 g/cm^ (9 
specimens). In Karagan red fox, this weight loading is 62 g/cm^ in 
males and 60 g/cm^ in females, and in red foxes from heavy snow 
regions, it is less: in Lapland red fox — ^27—30 g/cm^ (Nasimovich, 

302 1948) and in Pechora red fox — 28 g/cm^ (Teplova and Teplov, 
1947). Thanks to the great weight loading and relatively short limbs, 
the corsac fox deeply sinks into loose snow, and with snow cover 
depth of 15—20 cm, it moves with great difficulty. At that time, the 
corsac fox stays in burrows or uses trails and tracks of other 
animals and humans. In distinction from the red fox, the corsac fox 
does not have a "mouse-leap" — a unique jump on rodents found 
under the layer of loose snow. Therefore, with deep snow cover, 
corsac foxes seek out areas where the snow is blown off or com- 
pacted by wind, and they also are restricted to places of winter 
grazing by large herds of saigas and domestic livestock. In obtain- 
ing food from under the snow while winter grazing, these ungulates 
dig up and trampling it, help the corsac fox to move and to catch 
rodents (Skalon, 1952; A. A. Sludskii). In the presence of deep 
snow cover, hard-crusted or ice-covered ground, the animals suffer 
from hunger and leave that region. 

In places with tall grass cover (meadow grass) as, for instance, 
in Badkhyz (southern Turkmeniya), corsac foxes constantly use the 



441 



numerous trails beaten by goitered gazelle and kulan (V.G. Heptner). 
In winter, corsac foxes were observed at carrion during the day, 
and on windless sunny days, lying on the snow near a burrow 
(Sludskii, 1930; B.D. Abaturov). In Betpak-Dala, corsac foxes were 
met with in December around 10-11 in the morning, and during the 
afternoon, when they approached domiciles to feed on refuse. At 
that time, they were also active after dark. The same thing is 
known for southeastern Trans-Baikaliya and Mongolian People's 
Republic (Brom et al., 1948; Skalon, 1949, N.V. Nekipelov). In the 
northern part of the Mongolian Republic, the corsac fox in winter 
begins hunting at the end of the day, before evening and scurries 
about searching for food the entire first half of the night; it again 
hunts early in the morning, and very rarely during daytime. 

In summer, corsac foxes are also active not only at twilight and 
at night, but often hunt during the day, especially when they are 
feeding young (Nasimovich, 1951). In captivity, at air temperature 
from 3° to 10°C, corsac foxes were active at 1900, 2300 and 0200 
hours [Fig. 113]. With reduction of air temperature from 25 to 0°C, 
activity of the corsac fox decreased threefold (Kalabukhov and 
Poluzadova, 1946; Kalabukhov, 1950). 

Young animals, 1.5-2 months of age, assume a more diurnal 
mode of life. Thus, in Betpak-Dala, from 1 1 to 27 May, a litter of 
8 pups might be observed on the surface already at 5 in the morn- 
ing. Until 9 am, the corsac pups played near the burrow entrance, 

i 40 

CD 

I 28 
i 24 




9 и 



13 5 7 3 



301 



13 15 /7 19 21 23 
Hour of the day 

Fig. 1 13. Daily cycle of activity of corsac fox in an experiment at 3— 10°C 
(after Kalabukhov, 1950). 



442 

moving not more than 5 meters away. When it became hot, they 
hid in the burrow and by 1 1 am only individual animals remained on 
the surface, but in the course of the whole day 1 to 2 corsac pups 
usually were lying near the burrow, sometimes changing places. At 
the end of the day, the corsac pups came out from the burrows 
more frequently and remained on the surface for a longer time. By 
20:00 hours, the whole litter was already inside the burrow. Coming 
out of the burrow, the corsac pups looked around for a long time, 
moved slowly and often scratched; apparently, they were greatly 
disturbed by fleas. At sunset, they cautiously began to play. When 
threatened by danger, they broke off playing and stretching their 
necks, endeavored to see what had frightened them. When afraid, 
the pups barked, making a sound which may be given as "vyau- 
vyau". After quieting down, the corsac pups again returned to play 
which continued until the approach of full darkness. But even later, 
at 11, 12 and 1 at night, when the burrow was illuminated, the 
whole litter could be seen near it. 

When coming out of burrows, the corsac fox at first thrusts out 
its head only, then shows itself to the shoulder, and only after being 
sure that it is quiet all around, does it come out on the surface. 
After coming out, the animal stretches itself several times, looks 
around, and then sits down, or sometimes lies down, near the bur- 
row, continuing to observe its surroundings. Sensing danger, it sits 
and watches closely, slightly nodding its head. If the alarm is shown 
to be not false, it begins "barking" in that direction, continuing, 
however, to sit in its place. After quieting down, with the approach 
of twilight, it sets out on the hunt and already at 21:00 hours, the 
animal could be observed beating back and forth along the road for 
jerboas. 
303 Both in summer and in winter the corsac fox, lying near the 
burrow entrance, does not respond. to a human approaching it be- 
fore a distance of 30 to 50 meters; after that, it runs aside and does 
not run into the burrow. When followed, it runs to a neighboring 
burrow, and remains motionless. Thus, even injured animals behave 
in the same manner, but when escaping from a wolf or eagle, the 
corsac fox enters the first available burrow, even if inhabited by 
marmots. 

It is interesting that the corsac fox has the ability to remain 
motionless, feigning death. The animal is sometimes found thus in 
an excavated burrow. It lies completely without movement, relaxed 



443 

and with its eyes closed. Taken up in the hands, it remains in this 
state, completely lifeless. But, if it is put on the ground, it opens its 
eyes, jumps up and runs away. (Similar behavior is known for red 
foxes, jackals, striped hyaenas, ratels, raccoon dogs and especially 
opossums.) 

The corsac fox is somewhat timid and cautious. It permits 
humans to approach as near as 10 meters, and then runs off for 
100-200 m and remains motionless. It also permits a car as close 
as 5-6 meters. It is caught in traps more easily than the red fox. 
With abundant food, for example, at a time of massive death of 
ungulates from starvation, it sometimes remains with red foxes and 
even dogs near carrion at cattle burial grounds and dumps (Sludskii, 
1930). 

In summer, corsac foxes hunt alone: the animal slowly walks 
against the wind and on smelling or seeing prey, conceals itself by 
pressing to the ground and running from one cover to another. Not 
far from the intended victim, it jumps up and overtakes it in a few 
springs. Rarely, it runs at a slow gait, and startling its prey, for 
example, sagebrush vole, pursues its prey, and overtaking it, en- 
deavors to pin it with its front feet. Where jerboas are numerous, 
it watches for them by remaining motionless behind bushes along 
roads or paths, where these rodents run, at the approach of twilight 
or at sunrise. On the ground, it searches for nests and preying on 
nesting birds, eats the nestlings and eggs. It catches goslings when 
their parents lead them to the steppe to feed, and ducklings at the 
time the adults lead them from the nest to water or transfer them 
from one lake to another. The corsac fox hunts lizards by jumping 
on little shrubs under which they are hidden, and endeavoring to 
roust them into the open. 

In winter, it also searches for food singly. In the literature, 
there is some information stating that corsac foxes sometimes hunt 
in pairs or even in small packs. Actually, they gather as several 
individuals around large carrion, but approach it singly, following the 
tracks or trail trampled by other animals. 

In the search for scarce winter prey, the animal goes at a trot, 
stopping and sniffing. Scenting or hearing the squeak of an animal 
under the snow, it quickly digs the snow trying to catch the rodent. 
With snow cover deeper than 10-15 cm, or when it is compact or 
covered with a hard crust, the hunt of the corsac fox is usually 
unsuccessful. 



444 

Corsac fox possesses a good sense of smell. One can success- 
fully follow by tracks on the snow how it turned at a right angle to 
its path and 20-30 m to the side dug out from under the snow an 
old sheep leg, horse hoof or leather scrap (Brom, 1952). The corsac 
fox preys on willow ptarmigan and gray partridge night-roosting in 
the snow, black larks and snow buntings, and watch for white hares 
on their trails. In breaks in the reeds, the soil buried under the snow 
is not frozen, and it catches water voles by excavating their burrows. 

Corsac fox tames well, and can live in captivity for a long time. 

Seasonal migrations and transgression. The approach of 
cold, the entrance into hibernation of rodents, the 1офог of reptiles 
and insects, the migrations of a series of birds and the falling of 
snow force the corsac fox, in late autumn or the beginning of 
winter, to migrate to the south, but sometimes also in other direc- 
tions. Obtaining winter food, which is rare itself, becomes more 
difficult if snow cover is deep — ^more than 1 5 cm, or forms a hard 
304 snow crust or ice-covered ground. Abundance and availability of 
food depend also on other factors: severe frosts at the beginning of 
winter when there is still no snow, steppe fires, migration of ungu- 
lates, etc. All these phenomena often take place in the forest-steppe, 
steppe, and some in the northern deserts of Kazakhstan. 

In individual winters, the number of corsac foxes in the steppes 
increases from the tally of animals arriving from the forest-steppe, 
and with low numbers or difficulties in capturing rodents in the 
steppe, corsac foxes leave it for the semidesert. In the northern and 
central parts of its range, including the northern deserts, the sea- 
sonal migrations of these animals bear a more or less regular 
character and take place almost annually. In the beginning of win- 
ter, the majority of corsac foxes abandon summer places and search 
for sections richer in food (Polferov, 1896). In the steppes of north- 
em Kazakhstan, in the heavy snow winter of 1940/41, corsac foxes 
(and also wolves and red foxes), migrated en masse to the south- 
west to the Turgai river and were encountered in Priishmisk and 
other regions of Sevemo-Kazakhstansk district, where they had not 
been previously observed for many years (I.R. Zenkin). In the 
winter of 1942/43, in these same places, corsac foxes migrated to 
the north. One hunter there caught 56 animals in the season, at a 
time when, in the usual years, only 2—6 animals were caught (E.E. 
Chernov). In 1947 (in November), intensive emigration to the south 
of corsac foxes was observed there, connected with the low 



445 

number of mouse-like rodents in the places of their summer resi- 
dence. In the heavy snow winter of 1948/49, massive emigration of 
corsac foxes was recorded in Kustanai district. The animals went 
to the south, while from the steppes of Tselinograd district, some 
migrated to the south and others to the northeast, towards Om'sk 
and Pavlodar. As a result, corsac fox population decreased by 
about 40% (K.T. Popov). In 1950/51, in connection with the early 
formation of snow cover, which was very compact as a result of 
thawing and refreezing, forming a thick frozen ice-crust, from the 
middle of December massive emigration to the south was observed 
from Kustanai, Tselinograd and Karaganda districts. 

Such migrations of corsac foxes, having in different years vari- 
ous directions and intensity, depending upon weather and food 
abundance, were also observed in other parts of the range. By the 
middle of December, with the formation of a deep snow cover in 
the semidesert, the population of corsac foxes in Betpak-Dala in- 
creases as a result of migration from the north. At that time, their 
tracks are encountered in great numbers and the animals them- 
selves may often be observed. In the northern and central parts of 
the Betpak-Dala desert, with the formation of the snow cover, the 
corsac foxes begin to suffer from a great insufficiency of food, 
since of the rodents, only the red-tailed gerbil remains active and 
they rarely come out of their burrows at that time. Therefore, 
corsac foxes here in winter migrate after saiga herds, feeding on 
the carcasses of antelope killed by wolves or starvation. The ani- 
mals move at night. Each year in December, saigas move to the 
southeast for 4 to 5 days through the central region of the Betpak- 
Dala; in this short period or a bit later, the corsac foxes also 
migrate. By the end of December, the majority of corsac foxes 
have already followed the trail of the saigas southward to the Chu 
river valley. Those remaining suffer greatly fi"om hunger. There are 
no rodents, and, at this time, the remains of saiga carcasses killed 
by wolves, foxes and corsac foxes also disappear. The last animals 
remain without prey and are also forced to migrate, following the 
saigas. Moreover, in places of massive saiga concentrations, the 
snow is much beaten down and trampled by them, which facilitates 
the movement of corsac foxes and their capture of rodents. Many 
corsac foxes die from starvation and wolves. In winter in Trans- 
Baikaliya and the Mongolian Republic, corsac foxes follow the 
migrating herds of Mongolian gazelles (Brom, 1952 and others). 



446 

The cause of corsac fox migrations may also be steppe fires, 
which cover great areas. Migrations explain their appearance in the 
regions, where in summer they are absent or rarely met with. For 
instance, in a series of regions in Kokchetav district, they are found 
305 only in winter, appearing there at the end of December. The mag- 
nitude, and often also the distance of migration, depend upon the 
number of corsac foxes and the degree of change in conditions 
which induces migration (much snow, frozen crust, cold, steppe 
fires, etc.). 

Corsac foxes migrate also in the east — in Pri-Baikal, Trans- 
Baikaliya and the Mongolian People's Republic. Thus, in some 
years, they appear in the Selenga river valley in Kyakhtinsk district, 
penetrating there from the MPR (Fetisov, 1949; Surmach, 1957). In 
southeastern Trans-Baikaliya, during the time of winter migration, 
corsac fox sometimes transgress very far to the north and appear 
in forest-steppe regions (Aleksandrozavodsk region — Algachi, 
Man'kovo, Bokhto and Bazanov; Stretensk region — ^Kuenga river, 
left tributary of the Shilka; E. Pavlov, 1949; N.V. Nekipelov). In 
the last decades, mass appearance of corsac foxes in Trans-Baikaliya 
was noticed in 1946-1947 and in 1955. The main reason for the 
mass appearance of corsac foxes in Trans-Baikaliya was their 
migration from the northeastern part of the Mongolian Republic, 
where in 1942/1945 and in 1953 there was in places mass repro- 
duction of Brandt's vole, which later disappeared, that provoked the 
mass migration of corsac foxes northward (N.V. Nekipelov). In 
Trans-Baikaliya, corsac foxes migrate, moving out from regions 
where Brandt's vole and Daurian pika populations are reduced. 
In Trans-Baikaliya, the corsac fox migration is more rarely caused 
by deep snow cover (Cherkasov, 1884; Dorogostaiskii, 1935, 
Podarevskii, 1936). 

Extinction of rodents or other unfavorable conditions cause mass 
migration and settlement of corsac fox even beyond the range 
boundary. In this way, they sometimes populate new territories, but, 
as a rule, not for long. Thus, in separate years corsac foxes ap- 
peared in the middle Volga region, where they were hunted in 1930 
and 1933 in the Bol'sheglushitsk region. They also transgressed in 
1936 into the Bol'shechemigovsk region (Dobrokhotov, 1937). 

In the severe heavy-snow winter of 1955/56, corsac foxes 
appeared in the forest-steppe of the Volga region at Vol'sk city and 
Teplovka village, arriving, apparently, from the Trans- Volga steppes. 



447 

Transgressions hither of corsac foxes there are so rare that none 
of the old-timers remembers such an immigration (Kirikov, 1959). 
In the winter of 1943/44, corsac foxes were encountered in 
Oktyabr'sk, Bugul'minsk and Bavlinsk regions of the Tatar ASSR, 
where they had not previously been observed (Popov and Lukin, 
1949). In order to reach the regions mentioned, the animals must 
have traversed not less than 320 km in a straight line. In the 30' s 
of this century corsac foxes were caught in the Beloretsk region in 
Bashkiria, 280 km from the border of their permanent range. In the 
past century, corsac foxes appeared beyond the Ural range, in the 
Shadrinsk region, 240 km north of places where they usually occur 
(Sabaneev, 1874). In the winter of 1948/49, when the snowfall was 
heavy in Kazakhstan, corsac foxes were caught in Sorochinsk, 
Perevolotsk, Krasnokholmsk, Orenburgsk, Burtinsk, Troitsk and other 
regions of Orenburg district, 200 km and more to the north of the 
core habitat (Darkshevich, 1950). In 1952, there were many corsac 
foxes in Chelyabinsk district, whence they arrived from Kustanai 
and Severo-Kazakhstan districts. The same was also observed in 
western Siberia, where the corsac fox penetrated northward to 
Lebyazh'ir in Kurgan district, Tyukalinsk, Тага and Vengerov which 
are 50-200 km beyond the usual northern limits of their distribution 
(Shukhov, 1928; Kirikov, 1959, 1960). 

Massive emigration of corsac foxes from the steppes of 
Kazakhstan to the forest-steppe of western Siberia took place at 
the end of the lO's and start of the 20's and especially in the 40's 
of the current century. The First World War and the Civil War 
which followed it distracted many hunters from the fur harvest. 
The corsac fox catch was greatly reduced. At the same time, the 
fields were badly treated. It was not possible to harvest and thresh 
the ripened wheat; much arable land was abandoned, allowing mass 
reproduction of rodents in them. Under-harvest of corsac fox and 
the wealth of food reserves favored its reproduction and intensive 
colonization beyond the usual limits of the range. In 1917, corsac 
foxes appeared in great numbers in the Barabinsk steppe east of 
306 the Irtysh. Their number increased without pause until 1921. There- 
after, their population began to decrease, and by 1929 they 
completely disappeared (Sludskii, 1930). 

In the Altai territory, in the territory of the Kiprinsk region, 
corsac foxes were common up to 1916, then disappeared. In the 
winter of 1920/21 in this region, a mass appearance of this animal 



448 

was noted, moving to the north. By the end of the 20' s, corsac 
foxes disappeared there (Velizhanin, 1931). The same picture was 
also observed southward — in Semipalatinsk district (Selevin, 1930). 

Migration of corsac foxes also takes place in deserts, where it 
is associated with prolonged, continuous snow cover accompanied 
by mass death of ungulates. After the heavy-snow winter of 1949/ 
50, corsac foxes appeared in great numbers south of Zapadno- 
Kazakhstansk and Aktyubinsk districts and penetrated to the 
northern shore of the Caspian. Already in December 1948, many 
of these animals arrived from the north in the Syr-Dar'ya river 
valley (Sludskii, 1953). The corsac fox passed through from the 
north in significant numbers to Dzhambulsk and Alma-Atinsk dis- 
tricts and to the Chu valley. They appeared not only in the foothills 
but also low in the mountains. With the formation of deep snow 
cover, they began to return again to the sands of the Pri-Chuisk 
Muyunkum, but there was also much snow there, and the animals 
died of starvation. 

At the time of emigration from the northern deserts, corsac 
foxes crossed the ice of [Lake] Balkhash and appeared in southern 
Pri-Balkhash (winters of 1938/39; 1943/44, 1945/46, 1948/49 and 
especially 1950/51). In the foothills of the Dzhungarsk Alatau and 
in the Alakul' depression, corsac foxes appeared in 1938/39, 1947/ 
48 and 1948/49, i.e. in nearly the same winters as in southern Pri- 
Balkhash. In Zaisan depression, the corsac fox population sometimes 
increased on account of the animals emigrating hither from Xinjiang, 
which took place in November— December 1938. During the time of 
emigration of corsac foxes from northern-type deserts southward 
from the southern edge of the semideserts to the foothills of 
the Tien Shan, they traveled a straight-line distance of 400 to 
600 km. 

With food scarcity or difficulty in obtaining it, corsac foxes 
approach populated places, even large cities. Thus, in the severe 
winter of 1939/40, in Pavlodarsk district, they were encountered in 
great numbers near settlements, and regularly fed on meat-packing 
wastes near the city of Pavlodar. In the heavy-snow winter of 
1940/41, in Severo-Khazakhstansk district (Sovetsk region, near 
Barankul' settlement), corsac foxes appeared in such great num- 
bers that one hunter obtained more than 50 animals in the season. 
In the winter of 1948/49, in Karaganda district, corsac foxes and 
red foxes often appeared on the outskirts of settlements and within 



449 

the precincts of Karaganda, where they fed on rubbish; several 
emaciated animals were caught by hand. 

Therefore, for the corsac fox in its whole range, it is charac- 
terized by considerable displacement, govemed, first of all, by periodic 
food deficiency. In the forest-steppes, steppes, semideserts and 
deserts, the corsac fox performs migrations almost yearly. The 
direction of migration changes, but more often the animals go south- 
ward, rarely to the north or other directions. Distances traversed by 
corsac foxes during times of migration are not established, but the 
boundary of the range, in connection with emigration, "pulsates" 
within limits of 50-600 km. As a result of migration and emigration, 
the corsac fox population in one or another region may change as 
much as ten fold. 

Reproduction. The corsac fox is monogamous, the male also 
participating in nurturing the young. Apparently, having formed a 
pair, corsac foxes live together until one of the members dies. In 
the Moscow zoo, a male living with two females in one cage 
covered [mated with] only one of them. In the course of a year 
only one litter occurs. 

In Kazakhstan, in the steppe zone, rut in these carnivores starts 
in February; in the semidesert and desert zones — in January or the 
beginning of February; and in the steppes of southeastern Trans- 
Baikaliya, in February. At the time of rut, in the tracks orange-red 
307 spots are often met with, left in those places where the animal 
urinated. Two corsac foxes usually live near some burrows. The 
animals usually rut at night; at this time their peculiar barking is 
often heard. Several males run behind a female in heat for the first 
time, or for one left barren for some reason, and fight usually 
occurs between them. The female usually mates in the burrow. 

The duration of pregnancy is not accurately established, prob- 
ably equaling 52 days on the average. The earliest litter was found 
in Aktyubinsk district on 15 March 1953. In central Kazakhstan, at 
Lake Tengis, on 11 April 1957, a female was caught with 12 
embryos, the body length of which equaled 10 cm and the average 
weight, 30 g. In Kustanai district, the first litter was found on 25 
April, and between 5—15 May the young corsacs came out from the 
burrow. In Kokchetav, Tselinograd and Ural'sk districts, the young 
appeared at the burrows on 15-25 April. In Semipalatinsk and 
Vostochno-Kazakhstansk districts, they begin to emerge on 5—10 
April. In southeastern Trans-Baikaliya, a recently parturient female 



450 



was caught on 16 April 1961 (N.V. Nekipelov). Litters here appear 
at the end of April— beginning of May (Peshkov, 1954). In the 
Moscow zoo, pupping of the corsac fox was observed eight 
times — ^between 18 March and 10 May. From Table 37, one may 
judge litter size, and when young began to emerge from the burrows. 
On 10 June 1942, in the Zharminsk region of Semipalatinsk district, 
16 young of the same age were dug out from a burrow. 

Therefore, in Kazakhstan, there may occur in a corsac fox 
litter from 2 to 16 young, but more often 3-6 (average of 5 for 100 
samples). In the northeastern Cis-Caucasus, three young animals 
were found in a burrow (Heptner and Formozov, 1941). In the 
ephemeral deserts of Uzbekistan, the litter usually consists of from 

Table 37. The number of young in one litter of corsac 
fox in Kazakhstan 



N^ Number ^ 
N^f pups in 


3 


4 5 6 7 8 


9 10 


Total 
litters 


Average 
number 


NJitter 
Zones \^ 










in litter 


and districts N^ 




Number of occurrences 








True steppes: 
Kustanaisk, 1 


8 


6 2 7 2 2 


— 1 


29 


4.9 


Severo- 












Kazakhstansk, 












Pavlodarsk 












[districts] 












Dry steppes: 
Ural'sk — 


2 


3 5 2 — — 




12 


4.6 


Aktyubinsk 












Kokchetav and — 


8 


8 6 10 3 2 


1 — 


38 


5.0 


Tselinograd 












Semideserts: 












Karaganda — 


— 


— 4 1 5 1 


— — 


11 


6.3 


Vostochno- — 


2 


14—2 — 


— — 


9 


4.9 


Kazakhstansk 












and 












Semipalatinsk 












Deserts: 












Dzhambul — 


1 






1 


3.0 






Total 1 


21 


18 21 20 12 5 


1 1 


100 


5.0 


in %. 1 


21 


18 21 20 12 5 


1 1 


100 





451 

3-4 young, but many of them die while still in the burrow. The 
reason for juvenile mortality in the burrow period of life is believed 
by many hunters to be due to ants which attack the still-helpless 
animals. They also point to the possibility of diseases (Suleimanov, 
1939). In the steppes of southeastern Trans-Baikaliya, the litter 
308 contains from 5-7, and up to 10, animals (Cherkasov, 1884; Peshkov, 
1954). In the Mongolian Republic, in the Kerulen valley on 22 May 
1944, 7 young were found in an excavated cor sac fox burrow 
(Skalon, 1949), and in Choibalsan region, on 24 May — 1 1 pups half 
the size of adults (Kucheruk and Dunaeva, 1948). In the Mongolian 
Republic, the number of young animals in the litter varied from 2 
to 12 (Bannikov, 1954). In the Moscow zoo, in five litters there 
were 3, 4, 5, 6 and 7 young. 

It is not known whether fertility of corsac foxes increases 
significantly in years of food abundance as happens, for example, 
in arctic foxes, although the larger litters were found in the years 
with abundant sagebrush voles or Brandt's voles. 

Growth, development, and molt. Corsac pups are bom blind 
and with closed ear passages. They are covered with a light brown 
fluffy pelage. The tail is monocolored, while in the red fox its tip 
is always white. The tone of their fur becomes increasingly more 
and more yellow with age. 

Weight of newborns is 60-65 g; body length, 13.5-14.0 cm. 
They can see on day 14— 16. At 28 days of age, they begin to eat 
meat (E.A. Petrova). They grow rapidly. Thus, in central Kazakhstan 
young animals — ^females caught on 28 May 1960, weighed 245—262 g 
and of corsac pups from a litter captured on 29 May 1 960, males 
were 392-400 g and females, 378-405 g. In this same region, two 
young females caught on 31 July 1951 each weighed 1900 g al- 
ready, and a male caught on 3 August 1951—2050 g. In the 
Betpak-Dala desert, a young female caught on 1 1 May 1955 weighed 
1250 g, and a male caught on 17 May 1956-1200 g. Young females 
also captured there weighed: 20 September 1957—1800 g, and 13 
December 1957—2000 g. In Turkmeniya, on the southern edge of 
the Karakum, a young animal caught on 6 May was half the size 
of an adult. Weight of such an adult female is 2.3—2.5 kg, and of 
male, up to 2.6 kg. Consequently, at the beginning of May in 
Turkmeniya and the middle of this month in the northern deserts, 
young animals already have attained half the dimensions of adults. 
Farther north, they are smaller in May. However, already by the 



452 

beginning of August, the young weigh 1900-2000 g, i.e. shghtly less 
than adults, even in the semidesert. Apparently, the corsac pup 
completes its growth in 4—5 months. 

In the den burrow, the female lives excellently with the pups 
until they are 2 months of age. The male, though he participates in 
the raising of the young, apparently rests in another burrow or 
outside it, since in no case has he ever been found in any shelter 
with the pups (sample size is still small). When the pups approach 
full growth, the female sometimes lives with the male in one bur- 
row, while the young are in a neighboring one. 

In spring and summer, many fleas are found in occupied corsac 
fox burrows (Brom et al., 1948), which greatly bother the animals. 
Apparently, because of the parasites and dirt in the burrow, the 
female changes burrows 2—3 times, carrying the young with her. 

The young disperse early. Out of nine burrows with young, 
excavated in June— August, in only one burrow were two young 
revealed, while in the remainder, only one. Young and adult corsac 
foxes also change burrows often. 

In burrows excavated in November— December, 2—5 young ani- 
mals were often found (Dinnik, 1914; Sludskii, 1930, 1939; Heptner, 
1932). On Barsa-Kel'mes island, in December, three young males 
were caught in one burrow, and in a second — ^three males and one 
female (Sludskii, 1939). Apparently, with the onset of cold, the 
earlier dispersed young corsac foxes gather again in one burrow. 
It is not known whether they live in groups during the course of the 
whole winter, though there are references to litters of corsac foxes 
that separate only at the time of onset of rut (usually in February) 
or in spring. Assertions that several adult animals may live peace- 
fully in one burrow require verification. 
309 Duration of lactation is not established; a female caught in 
Betpak-Dala on 17 May in a burrow with three young that weighed 
1200 g already had no milk in her glands and her teats were dry. 

Females attain sexual maturity at the age of 9-10 months. In 28 
animals investigated at the age of 4 months, there were 53.4% 
males and 46.6% females. Under favorable conditions, the corsac 
fox becomes more fertile and therefore its numbers may quickly 
increase. 

It is accepted that the corsac fox molts two times per year — 
in spring and autumn. In spring, the winter pelage is completely 
replaced; it is not established whether the pelage is also completely 



453 

replaced in autumn or not. By the end of winter, the hairs lose their 
luster; gradually dulling, the guard hairs become dry and frayed and 
their ends bend. First of all, fur on the head and limbs is replaced, 
then fur on the nape and shoulders molts, later the anterior part of 
the back, side and thigh, then the posterior part of the back and, 
last, the rump and tail. Consequently, in spring the molt proceeds 
from head to tail. Spring molt lasts quite a long time. In the northern 
deserts (Barsa-Kel'mes Island), it is already initiated by February 
15 and lasts until the middle of May. In a nursing female caught on 
17 May 1956 in Betpak-Dala, the winter fur was nearly fully shed, 
and was retained only on the sides, rump and tail. Animals caught 
in this desert in August had summer fur, while those captured in the 
beginning of November well exhibited new fur, but it was still not 
mature. The growth of the [winter] pelage ends only at the end of 
November, just after severe cold makes its appearance. 

Consequently, growth of winter fur is in reverse order. First 
growth commences on the tail and rump, then on the thighs and 
back, ftirther on the nape and sides, and lastly on the paws, belly 
and head. 

Enemies, diseases, parasites, mortality and competitors. 
Everywhere within the boundaries of the Soviet Union and the 
Mongolian Republic the competitors of corsac fox as regards food 
are the wolf, red fox, and steppe polecat [Mustela eversmanni] 
and in the forest-steppe and steppe zones, also ermine and weasel; 
in the Barabinsk steppe and in Trans-Baikaliya — kolonok [M 
sibirica] and in the latter region the solongoi [M altaica] as well. In 
central Kazakhstan and in Trans-Baikaliya among the competitors of 
the corsac fox is also considered the manul [Felis manul] and 
in the deserts— marbled polecat [Vormela peregusna], steppe [F. s. 
libyca] and sand [F. margarita] cats, jackal and several others. 

In the spring— summer period, wolf feeds mainly on the same 
rodent species on which the corsac fox feeds, and is very common 
in some places. By comparing fur tanning of both species, one may 
see that in the Tselinograd district, for each corsac fox caught, 
there was 0.5 wolf and in Karganda district, 1.4 wolves. Red fox 
is an especially dangerous competitor for corsac fox. Throughout 
the whole year, it feeds on the same food as corsac fox. It is 
stronger than the corsac fox and more successfully catches rodents 
of moderate and large size; it is also better adapted to mousing in 
regions where the snow cover reaches 10 cm and more. Only in 



454 

the semideserts does the corsac fox, apparently, hunts more suc- 
cessfully than the red fox. The red, fox is ecologically more plastic 
than the corsac. In northern Kazakhstan, judging by fur tarming 
data during recent years, the red fox is numerically predominant 
over the corsac fox. Thus, in Kustanai district, 2000-3000 red foxes 
are caught and only 500-800 corsac foxes, in Severo-Kazakhstansk, 
2000-2400 red foxes and only 100-300 corsac foxes, and in 
Pavlodarsk, up to 2500 red foxes and 1000-1500 corsac foxes. 

In semideserts, before ploughing of virgin soil, corsac fox ob- 
viously predominated over red fox. In Tselinograd district, for 
example, in the 30' s of the current century, only some tens of red 
foxes and more than a thousand corsac foxes were caught. In the 
40's, already 400-500 red foxes were caught and 1000-1200 corsaa. 
In the 50 's, the population of red foxes greatly increased and to- 
gether with this the maximum catch; in a season, 2900 fox skins 
were prepared and at the same time 2400 corsac fox skins. There- 
fore, competition between these two species of carnivores rapidly 
310 increased. In northeastern Cis-Caucasus, in the season of 1924/25, 
one hunter took 15-30 corsac foxes and 10-20 red foxes (Heptner 
andFormozov, 1941). 

In the northern deserts, the red fox population was always 
much higher than the number of corsac foxes. According to tanning 
data, in Kzyl-Ordinsk district, 4 to 5 thousand red foxes are caught, 
maximum 8 thousand, and corsac foxes — ^200—400, up to 900 indi- 
viduals. In Alma-Atinsk district, the corsac foxes are still fewer: 
during the last ten years here, the maximum catch of red foxes 
reached 15 thousand, and corsac only 650 individuals. In the steppes 
of Trans-Baikaliya, corsac fox populations are predominant over 
red fox. 

Therefore, in virgin dry steppes and semideserts, corsac foxes 
predominate, and in deserts — red foxes. Ploughing of virgin lands 
leads to disappearance of corsac fox and growth in numbers of red 
fox, of which the Ukrainian steppes, and in recent years — ^also 
Kazakhstan, are obvious examples. Competition between red and 
corsac foxes for food becomes particularly intense in autumn and 
winter, when the majority of rodents enter hibernation, many birds 
migrate, and reptiles and insects are torpid. 

Throughout the range of the corsac fox, the steppe polecat is 
numerous, also a serious competitor of the corsac fox. In the 30 's 
of the present century in Kazakhstan, up to 500 thousand skins of 



455 

this carnivore were taken every year; i.e. one hundred times more 
than corsac foxes. The intensity of competitive relationship be- 
tween these animals can be partially judged by the frequency of 
their capture in traps at tarbagan marmot burrows. For a series of 
years in eastern Trans-Baikaliya, during the warm months, 105 
corsac foxes, 4 wolves, 9 weasels, 67 solongoi, 412 steppe polecats 
and 8 badgers were caught in traps set for marmots (Brom et al., 
1948). For every corsac fox, four polecats were obtained. Ermine, 
weasel, kolonok, solongoi, marbled polecat and wild cats, because 
of their relatively low number and small size, are not considered as 
serious competitors of the corsac fox. The corsac fox itself rarely 
attacks small carnivores. 

Among avian competitors of the corsac fox are manifested 
saker falcon (Falco cherrug), pallid and northern harrier {Circus 
macrourus and C. cyanus), steppe eagle {Aquila rapax) and 
rough-legged buzzard {Buteo lagopus). In the western range, the 
long-legged buzzard {Buteo rufinus) appears as a competitor of 
the corsac while in the east — in Trans-Baikaliya and the Mongolian 
Republic, it is the upland buzzard {Buteo hemilasius). The steppe 
eagle and, rarely, the upland buzzard are enemies of the corsac fox 
and with regard to saker falcon and common buzzard [Buteo buteo], 
this animal becomes a commensal (see above). 

The wolf is the greatest danger to the corsac fox. In the sec- 
ond half of winter, especially with much snow, wolves kill many 
corsac foxes (Velizhanin, 1931). Thus, in Tselinograd district, in the 
winter of 1947/48, in February, one hunter found six corsac foxes 
torn apart by wolves. In Semipalatinsk district, in March-April 1948/ 
49, a hunter found the remains of 10 corsac foxes eaten by wolves, 
and in the winter 1950/51, in the Kurchumsk region, in one day, 
remains of five corsac foxes eaten by wolves were found. Wolves 
attack mainly animals weakened by starvation, or eat their dead 
bodies. They rarely catch healthy animals if these are near a bur- 
row in which they can hide. A pursued corsac fox runs with a 
speed of 40-45 and up to 50 km/hour, but it can run thus for only 
15—20 minutes. The desert wolf runs with a speed of 40-45 and up 
to 55 km/hour, but is more enduring and in open places, can catch 
the pursued corsac fox, especially if a group of wolves is hunting. 
In warm times [of year], wolves attack corsac foxes rarely, but 
even more rarely catch them. Thus in Betpak-Dala, the remains of 
corsac foxes were found in 0.8% of the wolf feces collected in 



456 

May-June (864 samples). In 101 stomachs of wolves caught in 
November-December in the same desert, corsac fox remains were 
found in only one. Wolves also destroy corsac fox litters, tearing 
apart their burrows (P.T. Kryzhinovskii). However, the attempt to 
excavate the burrow does not always meet with success. 
31 1 The red fox also belongs among the enemies of the corsac fox. 
It destroys the corsac fox in winter when it suffers from food 
shortages. Cases of corsac foxes hunted by red foxes and car- 
casses of corsac consumed by red foxes are known in Kustanai, 
Severo-Kazakhstan, Pavlodar, Kokchetav and Tselinograd districts, 
i.e. in steppes and semideserts. Red fox attacks on corsac foxes 
were observed particularly often in the heavy-snow winters of 1947/ 
48, 1948/49, 1950/51 and 1953/54. As a rule, hunters found corsac 
foxes killed by red foxes in March. With absence of food, red foxes 
attack corsac foxes earlier — in November and December (D.N. 
Boiko). In summer, red foxes destroyed den burrows of corsac 
foxes and killed the young. This was observed in the Oktyabr'sk 
region of Severo-Kazakhstan district and the Kellerovsk region of 
Kokchetav district (S.G. Chmut and D.P. Fomenko). Decrease in 
corsac fox numbers in the semideserts and steppes of Kazakhstan 
was related to the rapid growth of the red fox population, crowding 
out and destroying the corsac foxes, in the same way as the wolf. 
With abundant food, red fox and corsac foxes coexist with one 
another. They were observed together at carcasses of fallen live- 
stock and shot saigas (Sludskii, 1930). 

The corsac fox suffers also from shepherds and stray dogs. 
During the driving of livestock to and from summer pastures, thou- 
sands of hungry dogs, which harass and kill all animals and birds 
which they encounter on their way, go with the herds and flocks 
(Sludskii, 1961). Particularly dangerous are the Asiatic borzois — the 
taza and vyborzka—yNbxoh catch up comparatively easily with corsac 
foxes. Dogs also destroy the burrows of this animal and wipe out 
the corsac pups. 

The golden eagle (Aquila chrysaetos) often attacks red foxes 
and with training successfully hunts corsac foxes. However, its 
influence on corsac fox populations is negligible. The steppe eagle 
[Aquila rapax] is a more serious enemy. Remains of corsac fox 
were met with several times near the nests of eagle and in their 
castings (Nasimovich, 1951). In some places, the steppe eagle is 
abundant and may kill a considerable number of corsac foxes. 



457 

Thus, in Kalmykia, in places where saigas give birth to young, 24 
to 40 eagles were observed in one field of view (Dal' et al., 1958). 
In rare cases, the upland buzzard (Buteo hemilasius) attacks corsac 
312 pups (Nasimovich, 1951). In Semipalatinsk district, at the end of 
winter, remains of three corsac foxes were found eaten by an 
eagle owl {Bubo bubo). In winter, in the forest-steppes, steppes 
and semideserts, the snowy owl {Nyctea scandiaca) may attack 
the corsac fox. 

Diseases of corsac fox remain nearly unstudied. It is known 
only that they suffer from rabies and carnivore distemper. In 1950 
and 1 95 1 , a distemper epizootic spread over the whole territory of 
Gur'ev and Zapadno-Kazakhstan districts, the western half of 
Aktyubinsk district, and adjacent Astrakhan, Volgograd, Saratov 
and Orenburg districts. In this vast territory, mass mortality was 
simultaneously observed in corsac fox, red fox, wolf, raccoon dog, 
steppe polecat and even dogs. Mortality of corsac foxes from 
carnivore distemper was noted in 1950 in the Chapaevsk region of 
this same area (Sludskii, 1954). Summer mortality of corsac foxes 
from an unknown infection occurred in places in the Kurchumsk 
region of Vostochno-Kazakhstan district at the end of April 1948, 
and in July— September 1951 also in the Leninsk region of Severo- 
Kazakhstan district, where separate hunters found up to three fallen 
animals. Rodent plague, induced by B. pestis* has not yet been 
established in corsac foxes, though several hundred corsac foxes 
were examined in the regions where plague is endemic. 

In a young corsac fox, caught in the Sovetsk region of Severo- 
Kazakhstan district on 27 July 1941, the neck and head were heavily 
infected with ticks (species not determined). The following four 
species of fleas were noted on animals from central Kazakhstan: 
Pulex irritans (large numbers), Ctenocephalides canis (rarely 
encountered), Ceratophyllus tristinus (rarely encountered) and 
Ctenopthalmus breviatus (rarely encountered). The first two flea 
species are specific parasites of carnivores and of the latter two, 
one is a parasite of ground squirrels and the other, of voles (Mikulin, 
1956). 

In southeastern Trans-Baikaliya, for a series of summers in May— 
Octrober 6, 413 fleas were collected from 105 corsac foxes between 
May and October, from which were identified (in % of total): P. 

*Now Yersinia pestis — Sci. Ed. 



458 

irritans L. — 70.2; Oropsylla silantiewi W. — 3.7; Chaetopsylla 
homoeus Rh. — 5.3; Frontopsylla elata luculenta J. et R. — ^2.0; 
Ctenophyllus hirticrus J. et R. — 11.7; Rhadinopsylla dahurica J. 
et R. — 3.0; Amphalius runatus J. et R. — 3.2; Ceratophyllus 
tesquorum sungaris J. — 0.1; Neopsylla bidentatiformes W. — 0.2; 
Neopsylla pleskei orientalis J. et Arg. — 0.05; Ophthalmopsylla 
praefecta J. et R. — 0.2; Frontopsylla wagneri J. — 0.07; 
Pectinoctenus pavlovskii J. — 0.07; Paraceras flabellum W. — 0.01; 
Archaeopsylla sinensis J. et R. — 0.07; Ceratophyllus calcarifer 
W. — 0.03; Amphipsylla vinogradovi J. — 0.01. 

Fleas not specific for carnivores and which parasitize mainly 
rodents in Trans-Baikaliya, occur on the corsac fox and are assign- 
able to 13 species and constituted 24.5% [of the total] (Brom et al., 
1948). In southeastern Trans-Baikaliya, the average number of fleas 
parasitizing a corsac fox were: in April 3.0, in May 35.3, in June 
10.9, in July 37.0, in August 32.4, in September 84.8 and in October 
188.9 individuals. In corsac fox burrows, there are many fleas. 
Since a great number of fleas parasitize the corsac fox, it is prob- 
able that it acts as a vector of fleas infected with plague (Brom et 
al., 1948). 

The mortality rate among corsac foxes from various factors is 
not known. Especially many of them die, apparently, from food 
shortage caused by the decreases in rodent populations or their 
unavailability due to the deep snow cover, or hard-crusted snow, or 
ice. 

In Kazakhstan, many corsac foxes died from starvation in the 
winters with prolonged snow cover of 1940/41, 1947/48, 1949/50, 
1950/51 and 1953/54. Thus, in January 1949, in the Dzhurunsk 
region of Aktyubinsk district, one hunter found 5 corsac and six red 
foxes dead from starvation. Starving animals allowed humans to 
approach them and they could be caught by hand. In southeastern 
Trans-Baikaliya, the deep snow cover is also fatal to corsac fox. 
313 Food shortage in winter in some years is manifested as the most 
important factor in corsac fox mortality, leading to a population 
decrease. 

Population dynamics. In one or another region, often very 
broad, the numbers of corsac foxes is subjected to great annual 
changes. In years with abundant food, their populations rapidly 
grow so long as unfavorable conditions of existence do not occur. 
Population growth may be very rapid. In 1930, on Barsa-Kel'mes 



459 




311 f ig 1 14. Adult corsac fox with throat crushed by red fox near its burrow. 
Badkhyz Preserve, southern Turkmeniya. 28 April 1962. Photograph by 

A.A. Sludskii. 

Island (Aral Sea), where the corsac fox was absent, 3 males and 
2 females escaped from cages. In 1933 and 1934, 77 corsac foxes — 
the whole population — were caught, which had increased 15 times 
in 3-4 years (Sludskii, 1939). 

Among the factors reducing the corsac population is a small 
number of rodents — ^voles and pikas in the northern half of the range 
and gerbils in the southern desert parts — ^as well as their unavailabil- 
ity due to the deep snow cover (deeper than 1 5 cm) or the formation 
of hard snow crust or ice. At this time, corsac foxes go hungry, 
become weakened and die in great numbers from starvation, en- 
emies (wolves, red foxes) and diseases. Reduction of the population 
usually is carried out in one season. 

In the northern Caucasus at the end of the nineteenth century, 
the corsac fox was considered a very. rare and vanishing species 
(Dinnik, 1914; Satunin, 1945). However, in 1921-1925, many of the 
animals were found there: One hunter caught 15—30 corsac foxes 
each season during that time (Heptner and Formozov, 1941). Dur- 
ing 1926 and especially 1927, the corsac fox population decreased 



460 

greatly and from 1932 to 1944 their skins were not prepared for 
tanning. From the 40' s the corsac population there gradually grew, 
and then rapidly increased to 1951. However, in 1952, their number 
again fell catastrophically (Vereshchagin, 1959). The cause of the 
drop in the corsac fox population in the Cis-Caucasus was strong 
dzhut [extensive periods of continuous snow cover] (1949/50, 1950/ 
51, 1952/53). 

In the steppes and semideserts of Kazakhstan, judging by the 
delivery of corsac fox skins to the Irbitsk fair, at the end of the 19th 
century, its number fluctuated 15-fold. In the period from 1880 to 
1895, it was lowest in 1880, 1890 (in 1881,* only 5 thousand skins 
were delivered to the fair and in 1 890, 2 thousand; both times were 
after the heavy dzhut of 1879/80 and 1888/89). The highest popu- 
lations occurred in places in 1883—1886; in 1884, 30 thousand skins 
were delivered to the fair, and in the following years, 25 and 15 
thousand skins. 

In the forest-steppes of western Siberia (Orenburg, Chelyabinsk, 
Om'sk and Novosibirsk districts) and in the northern steppe zone 
of Kazakhstan (Kustanai, Severo-Kazakhstan, Pavlodar, Semi- 
palatinsk and other districts), the corsac fox had nearly disappeared 
by 1910 (Korsakov, 1938), and reappeared again in 1916-1917. Its 
numbers reached a peak in 1923. 

In the dry steppes and semideserts (Ural'sk, Aktyubinsk, 
Tselinograd (the former Akmolinsk), Karaganda and other districts), 
the corsac fox began to appear in considerable numbers from 1919 
on. Before this year, the inhabitants of Tselinograd district did not 
know of it. It became especially numerous in 1921 and 1922, but 
in 1923, the population began to fall rapidly and by 1924, this animal 
had again become rare. 

Intensive reproduction of the corsac fox in the beginning of the 
20 's was attributed to several factors. In connection with the First 
World War, followed by the Civil War, the number of hunters 
8Ьаф1у decreased and the hunting economy nearly stopped around 
1920-1921, i.e. for 7—8 years. Many fields were abandoned and in 
them there was strong reproduction of rodents. In these same 
years in the steppes many livestock, especially horses died. All of 
this favored the rapid growth of the corsac fox population, and it 
quickly spread in all Kazakhstan and adjacent areas. In these years, 

*From context, probably should read 1880 — Sci. Ed. 



461 

in Kustanaisk, Severo-Kazakhstan and Pavlodar districts, individual 
hunters took 150 to 200 corsac fox skins per season. After the 
314 peak population (1921-1923), 225,445 skins were exported from the 
USSR in 1923-1924 and 1924-1925 — ^a record number for the entire 
period of existence of the trade (Kaplin, 1960). 

The rapid decline in the corsac fox population in the middle of 
the 20 's was the result of hunting of carnivores and the ploughing 
of fallow and virgin lands. After the end of the Civil War, many 
hunters returned to peaceful occupations and the steppes were 
flooded with agents of tens of tanning organizations which rapidly 
drove up the price of furs. At this time "everyone who was able 
went to the steppe with tools, weapons, shovels and barrels filled 
with water and ruthlessly proceeded to kill the little foxes in great 
numbers, digging and smoking them out of burrows in groups. This 
barbaric 'hunting' was practiced especially intensively at the begin- 
ning of winter with the first snowfall, when it was easy to find the 
corsac burrows through their tracks" (Selevin, 1925). This rapa- 
cious taking of corsac fox was also carried out in western Siberia 
where, moreover, the animals were pursued on horses with or 
without dogs, and poisoned with strychnine. As a result, at the end 
of the 20 's, the corsac fox in many places in Kazakhstan and 
western Siberia was "almost completely exterminated" (Zverev, 
1932). Thus, in Semipalatinsk governance in 1921/22, 1785 corsac 
skins were tanned; in 1922/23—8,012; in 1923/2^^19,344; in 1924/ 
25—2,312; in 1925/26—503; in 1926/27—345; and in 1927/28— only 
165 (Selevin, 1925, 1930; V.E., 1928). Overall, in four years, the 
corsac population had decreased 117 times. In Kazakhstan (as a 
whole) in 1924/25, 23,194 skins were tanned; in 1925/26—19,426; 
in 1926/27—7,934; in 1927/28—4,024; and in 1928/29—505 (Gross- 
man and Kogan, 1932). In the following years, its harvest was 
prohibited. 

Mass reproduction of the corsac fox, which was observed in 
the 20' s of the present century occurred more or less simulta- 
neously over a vast territory. A similar picture of reproduction in 
the 20 's and subsequent decrease in their population was observed 
in the Cis-Caucasus, lower Volga region, forest-steppes of western 
Siberia, and Kazakhstan. An analogous picture was observed in 
some places in Turkmeniya. Thus, at the beginning of the 20's on 
the plains at the foot of the Kopet-Dag, corsac fox almost com- 
pletely disappeared as a result of the application of strychnine. At 



462 

one poisoned carcass, as many as 13—14 corsac foxes were picked 
up in a night (V.G. Heptner). 

In the steppes and semideserts of Kazaldistan, the corsac fox 
again began to appear in the middle of the 30's. Thus, in Kustanai 
district in the northern regions, it appeared in 1935, but was still 
absent in the Semiozemii and other regions, where it appeared in 
small numbers (Naurzumsk preserve) in 1936. Only in the heavy- 
snow winter of 1940/41 did corsac foxes here become numerous, 
migrating from neighboring regions. A new population peak (judging 
by tanned skins) in northern Kazakhstan was in 1938/39. In Kzyl- 
Orda and Chimkent districts, the corsac fox was especially numerous 
in 1934 and 1935. 

The rise in the corsac fox population in the 30' s in Kazakhstan 
stopped in the winter 1939/40 when, in many areas, the animals 
suffered from a shaф food insufficiency. After this winter, the 
corsac population in the north of Kazakhstan, judging by tanned 
skins, decreased nearly two times. It continued decreasing through- 
out the whole of Kazakhstan after the severe heavy-snow winter 
of 1940/41. But in 1944, nearly everywhere in Kazakhstan, the 
number of corsac foxes again increased and they were particularly 
numerous in the dry steppes and semideserts of Kokchetav, 
Tselinograd and Karaganda districts. At the beginning of the 40's, 
the corsac fox population also began to increase in the forest- 
steppes of western Siberia, and it appeared in regions where they 
had long been absent (Makushinsk region of Kurgansk district, 
Sargatsk region of Omsk district and others). 

In February— March 1948, in the northern and central parts of 
Kazakhstan, corsac foxes died in great numbers from starvation 
(voles were few). The winter of 1948/49 had exceptionally deep 
315 snow, and again many animals died from hunger and enemies; the 
corsac population greatly decreased (Sludskii, 1953). In this winter, 
corsac foxes appeared in significant numbers in the steppes and 
forest-steppes of Orenburg district (Darkshevich, 1950). In the 50' s 
of this century in the northern steppes of Kazakhstan, an increase 
in the corsac fox population was observed in 1952, but they were 
especially numerous in 1956. In 1955/56, corsac foxes were also 
numerous in the steppes of Altai territory. After the severe winters 
of 1948/49 and especially 1950/51, corsac foxes appeared in rela- 
tively high numbers in the northern foothills of the Tien Shan in 
Kirgiziya and in southeastern Kazakhstan. 



463 

Subsequent significant decrease in corsac populations in the 
northern, central and southern regions (Betpak-Dala desert), 
induced by the absence of the sagebrush voles and other rodents, 
took place in the winter of 1959/60. In the 1960/61 season, there 
were few corsac foxes anywhere. In Kazakhstan as a whole, 
tanned corsac skins fluctuate approximately 15-fold. 

In the deserts of the southern type (ephemeral), judging by the 
data on tanned furs, the highest population of corsac fox between 
1950 and 1959 was in Uzbekistan in 1950 and 1956-1959, and in 
Turkmeniya during the time from 1948 to 1959 — in 1952 and 1956; 
and the lowest — in Uzbekistan in 1955 after the severe dzhut of 
1953-1954, and in Turkmeniya, in 1948 and 1953. In the southern 
deserts, the amplitude of population fluctuations of corsac fox is 
less than in the steppes and semideserts. In Uzbekistan, tanned 
skins of this animal varied only 1.5 annually, and in Turkmeniya, 3.3 
times. 

In the Daurian steppes of southeastern Trans-Baikaliya, the 
corsac fox population varied 18-fold over nine years (from 1940 to 
1948) (Yurgenson, 1955). Judging from tanning, the highest popu- 
lation of corsac foxes here was in 1946/47 and in 1955. The main 
reason for this population growth was migration from the Mongolian 
Republic. In separate areas of the steppes, the Trans-Baikaliya 
population of corsac foxes changed under the influence of local 
factors (N.V. Nekipelov): epizootics among tarbagan marmots, their 
extermination by the antiplague organization, mass reproduction of 
Brandt's voles and Daurian pikas favoring the growth of the food 
base of the corsac fox, causing increase in its population. 

On the whole, the number of corsac foxes in the territory of the 
USSR changes annually by 10 up to even 100 times. The most 
considerable change in corsac fox populations is found in the 
forest-steppes and steppes, where with unfavorable conditions, they 
completely disappear over vast areas for several years. Populations 
are more stable in the southern (ephemeral) deserts. Before a 
population peaks, it increases for several years. The high population 
is maintained for 1-2 years, and then decreases shaфly. True pe- 
riodicity is not observed in variations of corsac fox populations. 
During -the last 20 years, increases have followed after 2-4 years, 
usually after three years. The amount of tanned skins also fluctu- 
ates strongly. 
316 Field characteristics. This animal is very similar to the red 
fox, but is two times smaller in size. In contrast to red fox, the ftir 



464 





-"-^-A...vn"- 



о 




Fig. 115. Tracks of corsac fox. Left — ^track of Kazakhstan corsac fox, 
V.c. corsac L. (quite large form), on the snow in the vicinity of Dzhanybek 
(left bank of Volga), 20 October 1951. Right — track of Turkmeniyan corsac 
fox, V. с turcmenicus Ogn. (small form), on wet sand. Karakum desert 100 
km north of Ashkhabad, 21 January 1948. Sketch by A.N. Formozov, about 
2/3 of the natural size. 



color is uniform reddish gray. The tail is fluffy and long as in red 
fox and of one color, but does not have the white tip characteristic 
of the red fox. Ears are reddish-brown externally and not black as 
in red fox. 

Tracks proceed directly — in a line. It steps precisely, track in 
track. The distance between tracks is 6 to 8 cm. The size of the 
footprint in the dust is 4 x 3 cm. At the entrance to the burrow, it 
levels the ground. Near the burrow there are always many feces — 
small sausages made up of mouse-like rodent hairs. As a rule, it is 
met with singly. 

In the case of the appearance of danger, for example, when it 
notices a human near its burrow, the corsac "barks". The agitated 
call of this animal is similar to the bark of a red fox, but at the same 
time is somewhat reminiscent of the mewing of cat and sounds like 
"m-yaaa". A disturbed corsac fox also gives another call: "un! un! 
un!". The cry "m-yaaa!" can be also heard at night in winter, 



465 



Я^ЙЯ'Ш 




316 Fig. 116. Track of corsac fox on snow. Betpak-Dala desert. Kazakhstan. 
15 December 1956. Photograph A. A. Sludskii. 



when the animals feed on carrion, and also during the time of rut. 
Young animals at the age of 1.5 to 2 months, sensing danger, give 
a cry "vyau! vyau!" (A.S.) 

Practical SigniHcance 

Corsac fox is a fur-bearing animal, but its fur is of low value, 
because it has quite coarse guard hairs. Previously its fur was more 
important. Harvest of this animal has been practiced for a very long 
time — in Kazakhstan, for example, since the Bronze Age ("Andronov 
culture") and probably earlier. Later, corsac fox skins were used 
for a long time for payment of tribute. Thus, the nomads who 
inhabited Kazakhstan under Mongol subjugation in the 1 3th century 
were obliged to pay a tribute of furs, which included corsac fox 
skins. At the beginning of the 18th century the ТоЬоГ and Barabin 
Tatars delivered corsac fox skins as tribute (Kirikov, 1959, 1960). 
From the ends of the 17th and 18th centuries, corsac fox skins 
were a subject of brisk trade between Kazakhs and Russians. 



466 

"Kirghiz-Kaisaks caught corsac foxes in the Zayaitsk steppe with 
317 golden eagle and dogs, and overtaking them on horses, killed with 
whips a multitude and they exchanged the whole lot, from 40 to 50 
thousand, with Russians at Troitsk fort and in Orenburg. According 
to the Orenburg tariff, they were placed in value at 40 kopeks 
each.'^ The Kirghiz, in their trade and exchange between each 
other, used them almost like money and the price of their goods 
was estimated according to the price of corsac foxes" (Rychkov, 
1762; analogous information concerning the trade in corsac fox 
skins in Orenburg is contained in Pallas, 1773). At Yamyshev fort 
on the Irtysh and in Semipalatinsk, corsac fox skins were brought 
from Dzhungaria (which then included southeastern Kazakhstan) 
(Struve and Potanin, 1867). On the importance of corsac fox skins 
as barter, P.S. Pallas (1773) wrote that "As they (Kirghiz-Kaisaks; 
A.S.) had no money themselves, they evaluated everything with 
reference to horses and sheep, and instead of small coins, used 
wolf and corsac fox skins". Harvest of corsac foxes was also 
developed in the 18th century in the forest-steppes of western 
Siberia (Pallas, 1786). In Kazakhstan, corsac fox skins were of 
great economic importance in the 19th century also. From 1857 to 
1861*, from 5,213 to 14,546 corsac fox skins were exported annu- 
ally from the Kirghiz steppe through the Petropavlovsk, 
Presnogor'kovsk, Omsk and Koryakovsk railway districts alone 
(Krasovskii, 1868). 

In the 18th and 19th centuries, corsac fox trade was also de- 
veloped in Trans-Baikaliya, where in the middle of the past century 
corsac fox skins were bought "from the first hand" for 50-60 
silver kopeks per skin (Cherkasov, 1884) and exported to China, 
where they were in great demand. In the 60's of the 19th century, 
a total 40-50 thousand skins were bought each year in Russia 
(Kaplin, 1960). 

The size of the corsac fox catch in Kazakhstan and Middle 
Asia at the end of the 19th century can partly be judged by the 
delivery of skins to the Irbitsk fair (they were not sent to others). 
Sent (in thousands of skins) in 1881—5; 1884—30; 1885—25; 1886— 
15; 1887—5; 1888—15; 1889-^.5; 1890—6; 1891—2; 1892—6; 1893—6; 
1894 — 8; 1895 — 5. By the end of the 19th century, the corsac fox 
trade was mainly harvested in the Aralo-Caspian lowland in the 

^^At that time the price of a sheep was 30 to 40 kopeks. 
*In Russian original, 1961, a lapsus — Sci. Ed. 



467 

amount of "more than ten" thousand (Silant'ev, 1898; Turkin and 
Satunin, 1900). 

In 1923/24, in the USSR, 135.7 thousand skins of this animal 
were tanned; in 1925/26-^2.9; in 1927/28—12.2; in 1928/2^— 9,1 13 
thousand. Thereafter, in the majority of regions, harvest was pro- 
hibited for a series of years. In 1940, 12 thousand were again 
tanned; in 1946—20.4; in 1951—21.8; in 1955—23.8 and in 1956— 
28.3 (Kogan, 1931; Kaplin, 1960). 

In the beginning of the 20th century, about 1 5 thousand skins 
of corsac fox were imported annually ft^om Mongolia to Kalgan 
[China] (Bogolepov and Sobolev, 191 1), and in 1927—10,200; 1928— 
18,200; 1929—16,600; 1930— 3,800 and 1931— 24,000 (Blokhin, 1935). 
In this country, not less than 1 5—20% of the skins obtained remain 
in the hands of hunters (Bannikov, 1954). 

In the late 20' s and early 30' s of the current century, the world 
yield of corsac fox skins ranged fi"om 26 to 50 thousand, of which 
fewer than half were taken in the USSR, and more than 50% in the 
Mongolian Republic. Before 1938, the majority of corsac fox skins 
were exported from the USSR to England, USA, France and Ger- 
many. For 1923/24 and 1924/25, 225,445 skins were exported abroad 
for the sum of 1,783,107 rubles (Kaplin, 1960). The demand for 
corsac skins abroad increased greatly when in 1922/23, a method 
was discovered to dye them in various colors while preserving the 
silvery tips of the guard hairs. 

In the 30's, as a result of the severe drop in populations of this 
animal, tanning of its skins greatly decreased, and together with this 
exports declined. After the end of the Second World War, the 
reduction in demand for long-haired furs influenced the price of 
corsac skins — ^they were greatly reduced, as a result of which their 
318 export in 1951—1958 was no longer profitable. On the eve of First 
World War, a corsac fox skin brought 1.2 dollars on the external 
market, in 1923/25 — 3.85 and in 1943—5.80 for a skin of first sort 
of the Petropavlovsk type. In the following years, the price de- 
clined, and then began to rise again (Kaplin, 1960). 

Earlier, in Kazakhstan and western Siberia, corsac foxes were 
caught in jaw traps, coursed with borzoi dogs and golden eagles, 
chased down on horseback and killed with whips and soila (long 
sticks), dug and smoked out of the burrows with smoke from burn- 
ing sulfur or grass, or drowned with water. Lastly, they were poisoned 
with strychnine, nuxvomica, and other local poisons. In Trans- 



468 

Baikaliya, they were caught with jaw traps, '"bashmalc", cherkan* 
and other traps which were set at the entrance to the burrow. At 
the present time, the main mass of the animals is caught with No. 
3 and No. 5 jaw traps and they are rarely coursed with dogs. 

The corsac fox is not a significant threat to small-animal hus- 
bandry. It usually causes little harm to the hunting industry, especially 
in the period of mass reproduction of mouse-like rodents, on which 
it mainly feeds. In those years with low populations of small ro- 
dents, does it sometimes attack large birds and hares. When 
populations are high, it may hunt the yellow ground squirrels and 
marmots; therefore in regions where these animals live, the number 
of corsac fox ought to be limited. In some regions, the corsac fox 
damages the hunting industry, eating animals that have fallen into 
traps, or dragging them off. Thus, in the Esil'sk region of Severo- 
Kazakhstan district, in the season of 1936/37, corsac foxes ruined 
300 polecats which were taken in traps (Sludskii, 1939). 

In a series of republics and districts, the corsac fox is protected 
as a valuable fur-bearer. Thus, it is completely forbidden to dig 
them out of burrows, smoke them out or drowned them. In the 
majority of republics, hunting this animal is only permitted in the 
season when it has prime fur — ^usually in November-March. In other 
times of year, the season is closed. 

The Standard of Fur Raw Materials divides the untanned skin 
of corsac fox into four types according to the region where they 
were obtained and quality of the fiir: Petropavlovsk, Kazakhstan, 
Astrakhan and Tashkent, of which the first type is the best and the 
latter is the poorest. 

In periods when populations of this animal fell catastrophically, 
harvest was completely prohibited for many years over large ter- 
ritories. For example, in the majority of the regions of Kazakhstan, 
hunting of corsac foxes was prohibited from 1928 to 1938. 

Hunting was permitted only in seven regions where a ground 
squirrel-gerbil industry existed (Irgizsk, Turgaisk, Karsakpaisk and 
others). Harvest of corsac fox was prohibited until 1939-1942 in 
districts north of Kazakhstan, for example, in Orenburg district 
from 1932 to 1939. This prohibition together with other factors 
favored the restoration of the corsac fox population. In the gerbil, 
marmot and other game industries, corsac fox populations ought to 
be restricted, without, however, destroying them completely. (A.S.) 

* Local names of different trap types — Sci. Ed. 



469 
Fox, Vixen, Red Fox'^ 
Vulpes vulpes Linnaeus, 1758 

1758. Canis vulpes. Linnaeus. Syst. Naturae, ed. X, 1, p. 40. 

Uppsala, Sweden. 
1758. Canis alopex. Linnaeus. Ibidem, p. 40. Uppsala, Sweden. 
1777. Canis caragan* Erxleben. Syst. Regni Anim. Mammalia, 

p. 566. Area of the lower Irgiz and Turgai [rivers] north of 

the Aral Sea.'^ 
319 1789. Canis crucigera. Bechstein. Gemeinn. Nat. Deutschlands, 

1, p. 250. Thuringia. 
1811. Canis melanotus. Pallas. Zoographia Rosso-Asiat., 1, p. 44. 

Kazakhstan south of Orenburg (V.H.). 
1816. Vulpes vulgaris. Oken. Lehrb. d. Naturg., 3, Th. 2, p. 1034. 
1836. Canis vulpes montana. Pearson. Joum. Asiat. Soc. Bengal, 

5, p. 313. Himalaya. 
1843. Vulpes flavescens . Gray. Ann. Mag. Nat. Hist., 11, p. 118. 

Northern Iran. 
1854. Vulpes pusilla. Blyth. Ibidem, p. 729. Salt range, Punjab. 
1854. Vulpes leucopus. Blyth. Ibidem, p. 729. Mal'tan (Multan), 

Punjab. 
1875. {Canis vulpes) var. beringiana. Middendorf. Ubersicht d. 

Natur Nord- und Ost-Sibiriens, 4, Th.2, 1 Lief. p. 990. Asi- 
atic coast of Bering Strait (V.H.) ("an den West-als an den 

Ost-Kusten des Berings-Armes"). 



^^Hunters often call the male "lisovin". 

^''Erxleben {Mil) stated that this form inhabited the "Kalmytsk and Kirgiz 
deserts", i.e. steppes, referring to it in two places in Vol. 1 of Pallas "Reise" 
(1811). The first reference (pp. 199-200) regarding the area south of the Samara 
319 river, is not precisely determined and may not require attention. The second (p. 
234) refers to the area south of Orenburg where "the Kirgizes bring the steppe 
foxes (karagan) which do not differ in color from wolves" to be sold in Orenburg 
(and hence the diagnosis of Erxleben — "colore griseo-lupino"). The above men- 
tioned type locality of the subspecies represents a more precise definition to 
references based on the usually cited type localities ("Kalmytsk and Kirghizian 
steppes", "Kirghizian steppes", "Kazakhstan") which are extremely broad. 
Karagans are, apparently, quite characteristic of the suggested limits of the type 
locality. Erxleben (1777) considered the karagan as "a doubtful animal" (animal 
subobscurum). Later, Pallas (1811) considered it to have the status of a separate 
species differentiated ft"om the red fox. 

*Sometimes spelled karagan — Sci. Ed. 



470 

1902. Vulpes vulpes splendens. Thomas. Ann. Mag. Nat. Hist., 
10, p. 489. Astrabad, northern Iran. 

1903. Vulpes anadyrensis. J. Allen. Bull. Am. Mus. Nat. Hist., 
19, p. 167. Markovo on the Anadyr [river]. 

1905. Vulpes alpherakyi. Satunin. Izv. Kavkazsk. Muzeya, 2, 
No. 1, p. 146. Geok-Tapa. Areshsk Co., Elizavetopol'sk Gub. 
[governance]. 

1905. Vulpes kurdistanica. Satunin. Ibidem, p. 48. Gel'sk depres- 
sion, form. Karssk district, Turkey. 

1907. Vulpes vulpes tarimensis. Matschie. Wiss. Ergebn. Filchner 
Exped. nach China, 10, T.l, p. 164. Tarim. 

1907. Vulpes vulpes tschiliensis. Matschie. Ibidem, p. 169. Beijing. 

1911. Vulpes vulpes septentrionalis. Brass. Aus. dem Reiche 
der Pelze, p. 454. Norway. 

1911. Vulpes kamtschadensis. Brass. Ibidem, p. 456. Kamchatka. 

1912. Vulpes vulpes flavescens var. cinerascens. Birula. 
Ezhegodn. Zool. Muzeya Akad. nauk, 17, p. 254. Eastern 
Khorassan, Iran. 

1914. Vulpes alopex var. caucasica. Dinnik. Zapiski Kavk. otd. 

Russk. geogr. Obshch., 26. Zveri Kavkaza [Animals of the 

Caucasus], 2, p. 449. Vladikavkaz (after Ognev, 1926). 
1914. Vulpes vulpes krymea montana*. Brauner. Zapiski 

Novoross. Obshch. estestvoisp., 40, p. 15. Montane Crimea. 
1914. Vulpes vulpes crymensis. Brauner. Ibidem, p. 36. Montane 

Crimea. 
1914. Vulpes vulpes crucigera n. stepensis. Brauner. Ibidem, 

p. 15. Steppes between the Dnepr and Don [rivers], more 

exactly — ^Askaniya-Nova (V.H.).'^ 
1922. Vulpes alopex var. kamtschatica. Dybowski. Archiw Tow. 

Nauk. Lwowe, 1, p. 350. Kamchatka. Nomen nudum. 
1922. Vulpes alopex var. sibiricus. Dybowski. Ibidem, p. 350. 

Nomen nudum. 
1922. Vulpes alopex var. ussuriensis. Dybowski. Ibidem, p. 350. 

Nomen nudum. 
320 1922. Vulpes vulpes jacutensis. Ognev, Biologicheskie izvestiya, 

1, p. 116. Vic. of Yakutsk. 



*Sometimes spelled krimeamontana — Sci. Ed. 
^^Vicinity of Kherson, after Ognev, 1 926. 



471 

1923. Vulpes hull. Sowerby.* Naturalist in Manchuria, 2, p. 44. 
Manchuria. 

1924. Vulpes vulpes crucigera diluta. Ognev et Vorobiev. Fauna 
pozvonochn. Voron. gub. p. 102, Rocky steppe. Bobrovsk 
u[ezd] [county], Voronezhsk. Gub. [governance]. 

1924. Vulpes vulpes schrenkii. Kishida. Mon. Jap. Mammals, 

p. 41. Sakhalin. 
1924. Vulpes vulpes splendidissima. Kishida. Ibidem, p. 47. 

Northern and middle Kuril Islands. 
1926. Vulpes vulpes alticola Ognev. Uchen. zap. Severo-kavk. 

inst. kraevedeniya, 1, p. 52. Lake Gokcha (Sevan), Armenia. 
1926^^. Vulpes vulpes karagan natio fergane ns is. Ognev. Annales 

Musei Nat. Hungarici, 23, p. 222. Osh, Fergana. 
1926. Vulpes vulpes karagan natio pamirensis Ognev. Ibidem, 

p. 222. Near Pamir Post, Pamirs. 
1926. Vulpes vulpes ochroxantha Ognev. Ibidem, p. 225. Aksai, 

Tien Shan. 
1926. Vulpes vulpes tobolica Ognev. Ibidem, p. 227. Obdorsk, 

lower Ob' [river]. 
1926. Vulpes dolichocrania Ognev. Ibidem, p. 232. Sidemi in 

southern Ussuri territory. 
1929. Vulpes vulpes kiyomassai Kishida et Mori. Lansania, I, 

p. 82. Northeastern Korea. 
1931. Vulpes vulpes daurica Ognev. Zveri Vost. Evropy i Sev. 

Asii, 2, p. 331. Kharagoi, 45 km west of Troitskosavsk 

(Kyakhta), Trans-Baikaliya (V.H.). 

Diagnosis 

Measurements large — ^body length more than 65 cm, greatest length 
of skull more than 115 mm. General color from light-reddish to 
grayish-ochre, brown; dorsal side of ear black, tip of tail white 
(V.H.). 



*In original text, misspelled "Sowerdy" — Sci. Ed. 
^^Issued in the beginning of 1927 (April; V.H.). 



472 

Description 

The red fox is a very well-formed, beautiful animal with an elon- 
gated body on relatively short limbs. The tail is long (longer than 
half the body length), fluffy, and in the standing animal reaches the 
ground. The muzzle is quite long and pointed. 

In its short summer coat, the red fox appears lean with rela- 
tively long limbs, with a thin elongated body, large head and large 
erect ears broad at the base and pointed at the tips. Northern foxes, 
in their fluffy winter coat, look stocky with short limbs, massive 
body and a proportionally moderate-size head. The ears, the basal 
portion of which are hidden in the fur, are more proportional [ap- 
pear smaller], the tail is especially long and fluffy, its overall length 
with hairs is nearly equal to the body length. ^*^ Southern forms of 
red foxes, in which the winter fur is relatively short, appear in 
winter as longer legged and "leaner", their ears are larger — ^taller, 
and broader at the base. 

Pupil of the eye is oval, and vertically oriented. There are three 
pairs of teats. 

The winter pelage of the red fox is dense, soft, silky and rela- 
tively long. However, in connection with its very wide distribution 
and its occurrence under extremely different climatic conditions — 
from tundra to hot deserts — the geographical variation in this character 
is very great. In the northern forms, the fur is very long, dense and 
321 fluffy, while in the southern— much shorter, sparser and coarser 
(see section on "Geographic Variation"). 

Still more variable is color of the winter fur of the red fox.^' 
This [variation] is especially complicated and great because, on a 
level with the well expressed geographic variation is wide individual 
variation. Besides form and amplitude of the latter, variation also is 
observed in types of moфhisms. The picture is further complicated 
in that in various subspecies, not only is the extent of natural vari- 
ability not the same, but also various color phases and types are 
encountered in various percentage ratios or are completely absent. 
For the latitude of variation in color and quality of fur, this species 



^"Hunters very neatly call it "tube". 

^^Color variation in summer fur was not studied, but judged by everything, 
it is less. 



473 




Fig. 117. Red fox, Vulpes vulpes L. Sketch by A.N. Komarov. 

is divided into 41 sorts, most of which are in our animals. Wit